THE METAZOAN PARASITE FAUNA OF LOONS (AVES: GAVIIFORMES), ITS RELATIONSHIP TO THE BIRDS' EVOLUTIONARY HISTORY AND BIOLOGY, AND A COMPARISON WITH THE PARASITE FAUNA OF GREBES
BY ROBERT W. STORER
MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITYOFMICHIGAN, NO. 191
Ann Arbor, June 18,2002 ISSN 0076-8405 l'he r~~t)licatiorrsol'the MLIS~LIIT~of Zoology, 'Il'hr Ilnivrrsity of Michigan, consist pr-ilnal-ilyof two scl.ics-the Mi,cc.ollcrr~~ou.sI'c~/)~r.salrtl tlie Occ.ci~ionc~lPi~l)lic.nlior~.\.Both series were loulrtlctl Ily 1)n KI-yant Walker; MI-.Bl-adsllaw H. Swales, ant1 Dr. W. W. Newcornb. Occasiollally the Museum publislrcs contl-i- butions outside of these series; I~egilltririgiri 1990 these ;lt-r titled S/)rc.ic~lI'u1~lic.nlion.s and 211-c rluln- bered. All srrl)lr~irtrdtrlatruscripts rrceivc external I-eview. Tlie Mi,\c.(~lla.nc~o~i.\IA~rDlic.ntion.s, which iricludc rnonogr;~pJricstrtdies, papers on lieltl alrd tli11sc111n tecllniqucs, and other contriblrtions not withi11 the scopc of tlic 0c.c.ntionnlI'np~\, are r)ublish(.tl sept- mtcly. it is ~iotilltended chiit they bt. grouped illto volurllcs. Each ~iurnbcr-has a title page and, wheli 11ecrss;u-y, a tablv ol'c.olrterrts. 'l'he Oc.c.c~sionn1I'(L/)(Ts, pllblicatiotl of' wllicli was beg1111 in 1913, scrvc ;IS a metlium 101- 01-igin;~l studies basetl principally upon tlle collections ill tlrc Musc~lrn.They at-e issiled srl)aratc,ly. M'lreti a sufficient n11mbt.l-of pages has I)?rn pri11tc.tl to rrlakc a volurric, it title pagc, table of contc.llts, :111tl an index il1.e slrpplirtl to lihr.;u.ies and illdividuals on the mailing list for tlie set-ies. A cornplctc. list of l,rlblicatiotrs on Bit-ds, Fishes, I~isccts,Mamlnals, Mollusks, 12cptilcs i111tlAtllpllil>- ians, ;lnd othcr topics is available. Address inquit-irs to the Dit.ccto1; M~lsclrrnof%oolo~y, '1-lie LJnivt*l-- sity of Michigall, All11 AI-hor, Michigan 48 105)-1079.
All,rrt, J.S. 2001. Species diversity alltl ~)liyloge~icti~systcln;rtics of' Anlet-ican knif'tl'ishrs (C;ylnnotili)rrnc.s, 'lklcostei). Misc.. PII~~.MIIS. Zool. Uriiv. Mielrigall, I!)O:I-127, 50 figs. N~~ssbat~tm,K.A. & <;:I. Raxwor-thy. 2000. Sysrcrna~icrevision of the genus I'UI-OO(~UM<;iilrthel. (Ke11- tilia: S(lllill~lat;~:Gekkorlidae), with thr descriptioll of' five licw species. Misc. 1'11111. Is!). 26 pp., 12 figs., 7 tables. Stover, li. W. 2000. l'hc mcti~zo;~~~pal-asite liut~aof'gt-rl~rs (Aves: Poilicipcttifi)~.~ilcs)ir~itl its rclirtion- ship lo tlrc birtls' biology. Misc. Prtbl. 188. 90 PI).,9 ligs., 7 tables. Nussbar~~n,1i.A. & M.E. PSrcntlcl: 1998. Revision of the African ciiecili;~ngetllls Srltislom~rlo/~~trn1':~-krt- (Amphil~iit:<:ylilrrol~hioria: (:acciliidac). Misc. I'ubl. 187. 35 pp., 15 figs., 15 tahlcs, 2 colol- plittcs. Nussba~uir,li.A., (::I. R'rxwot-thy & 0. Pr-o~lk.19!)8. The glrost geckos of' M;iti>igasc.:i~.:a lil~.tllu~. rcvi- siorl or tlle M;~l;~gasyIeal~toetl geckos (Reptilia, Squ;~~rlata,(kkkonitlae). Misc. Puhl. 186. 26 pp., 25 figs., 5 tal)lcs.
Not-I-is,S.M. 2001. Osteolog of tl~cso11tllwcstel.rl tl;+rtcl.s, I://~ro.\totncc(Oligoc.~/)hnlii.s) (Trlrostri. Percidi~e)-withcompi~risorl to other North Amet-ican percid fislres. Occ. pap. 733. 44 pp., 18 figs. Ng, H.H. & W:[. 1Ciint)otIi. 2001. i\ rcvicw of' tllc sisoritl catfish gcnlls 0roogLa11i.s(Silul-ifi)rmcs: Sisoritlae) with descriptions 01' li)lrr new spec.ies. 0cc. Pap. 732. 31 pp., I3 ligs., 3 cables. <:ollcr tc, R.R. 200 1. Oj)nsn'r~usekic-/~ro.stotnu.s, a new toatlfisli ('kl(:ostci: Bi~trachoididat)horn the wcstet.11 Oa~-it)l)ea~iSea arid soutllern (:uIf of Mexico. Occ. Pap. 73 1. I (ipp., 5 figs. Fink, W. I,. & A. M;~chado-Alliso11.2001. S(7m,snlrnu~.the~stntus, a tlcw spccics otpi~~nhafi.orn Krafil, with corrlliictits or1 S(v-rcl.\nImi~.cnllrruri and Srrrcrstrltnuc con~,t~r(~s.sus('l'elcostci: (:llaracifo~~~iics).Occ. Pilp. 7:30. 18 pp., 16 figs. GraP, U.1,. 2000. 'I'he Ethel-ioidei~I-rvisitetl: a pliylogcrictic iillalysis otllyr-iitl relationships (Mollllscil: Rivalvii~: P;~lcohctct-otlolrta:IJnionoida). Occ. I'ap. 729. 21 pp., 2 figs., 3 t;ll,les. Rarnanatrlaliji~to,J. R., li. A. Nussbautn & (:..j. Kaxwor-thy. 1999. A new species of' Mnh,~ry(~Fitzinger (Sq~la~nat;~:Scincitlirr: 1.ygosorninae) fi-om nol-thern Madagascan Occ. Pap. 528. 22 pp., 5 figs., '3 titblcs. Sp;rrks,.I. S. & P. N. Kcinthal. I!)!K). I'c/rc.lt-oj)lu.\ tnaromc~ndin,a new cichlid fish SI-or11the not-thwcst 01 Madagascar: Occ. l'irp. 727. 18 pp., 5 figs., 3 tahlcs.
TI3E KE(;ENTS OF THE LJNlVEKSlTY
H.,joscph Wl~irc,1.x c~/]irio I);ll~irl1). Horllir~g,(;rand Havrrr Andrra Fisc11c.r Nc\\~rnal),ht111 iZrl>or D;ivitl A. Rfiurtlo~~,All11 Arl,ol- Olivia P. Maynard, (hodrich S. Mart~n7:,iylor, C;rossc I'oi~rccFar-ms I.;ulrcl~cr8. Lleitcl~,Kloo~r~licld I lills Kcbecc;~McGowari, An11 Arbor ICatherine E. White, Ann Arl,or The Metazoan Parasite Fauna of Loons (Aves: Gaviiformes), Its Relationship to the Birds' Evolutionary History and Biology, and a Comparison with the Parasite Fauna of Grebes
Robert W. Storer
Bird Division Museum of Zoology The University of Michigan Ann Arbor, Michigan 48 109-1079, USA
MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 191
Ann Arbor, June 18, 2002 Ann ilrboi-,June 18, 2002
The life cycle of the digenean, Micr@hnlllrs nicolli. (a) The adult parasite, whicl~reaches a Inearl length oS0.54 nlm, inhabits the small intestine of the definitive host, a Co~nrnonLoon, (;n.r/ic~irn,nur (b).The minute (0.02 mm) egg (c) is passed illto the water where it hatches as a iniricidiurn (d) which penetrates the first intermediate host a snail, B%tt%umc~lt(~r1,aturn (e) in which cercai-ia (I') are produced. These penetr-ate the second intermediate host, a blue crab, Cn1li.necle.s sn;l,%tl~is(g). The definitive loon host becomes infectcd by eating by the crab. Infections of this parasite in the Common Loon occur most frequently when the bii-ds are unable to catch their usual, kxer- moving prcy, lishes, and it has been coninionly Pound in ailing loons on the wintering grounds oft'Floi.ida. Microphallids like this one show marked specilicity in their two intermediate hosts (first a snail, and then a crustacean), but are generalists i11 their definitive hosts. Original drawing by,John Megahan from sources listed in the ackrlowledglner~ts(p.32).
Slo7i.r; X. W 2002. 7Xr motnzoon fjn.rtc,sil(>Jiczrlra ojloons (Aucs: (;ar,jj/Orm,r~.),its ~rlnlon,,shiplo lhfj hids' c.-i,olution.crry hi.slo,rj~an,d biology, and (1 co,rnf)nrisonru%tlz th~pornsZt~/nulzn ofgrebes. iV1zsc. PubL. MILS.Zool. Ulziv. Michigan,, 191: zv+l- 44, 4Jg:s, 7 tcrblcs. Tllc data base of this work parallels that of Storcr (2000) on the grebes and contains lists of the multicellli1a1-parasites known to parasitize loons and a list of the known species of prey taken by each species of loon. The forrner includes information on w1lt.1-e in the bird the parasites are found, the distribution of the parasite species by continents, the degree of'host specificity, and life cycles (whether in fresh or salt water), and lists of' kilown intermediate and paratenic hosts. These data sets are used to sllow how the parasite bunas are related to the l~iologyof the birds and their evolutionary histories. The known species of hclminths of loons incltide 47 digeneans, 22 cestodes, 14 acanthocephalans, and 15 nematodes, niost ofwhich have aquatic life cycles. No families or suhfilrnilies;-~ild only 2 genera and 23 species of helrninths are considered loon specialists. Several [actors appear to have contributed to the greater numbers oS genera and species of parasites in grebes than loons. Most loons spend the breeding season 011 bodies of oligotrophic watel; which have fewer species and nunlbci-s of potential prey than the eutropl~icwaters where most grebes nest. All species ol'loons winter on salt water, whereas inally grebes are resident on fresh waters, althol~ghthose nesting in regions where tlie fresh waters f'reeze in winter spend that season on salt waters. Much of the difference can also be attributed to the greater number of'genera (7 versus 1 ) and species (21 versus 5) in the two groups of the birds, and the wider distribution (nearly cosmopolita~lin the grebes 71ersu.sholarctic in the loons). The diff'erence is also consistent with the grebes' 91-eaterage and degree of parasite host specificity. Otller tjctors aLfecting differences in the two l~elmintl~faunas are a preslnned marine origin for the loons in contrast with a fi-esh-waterone [or the grebes, and the greater size ailcl hence greater spccd of' loons underwatcr. The smaller size of grcbcs for wllich a greater number of prey species ol' optimal size is presumably available, the grebes' breeding on eutrophic bodies of water in which a greater variety ofprey species (and l~enccgreater number of species of hosts for parasites) is available, and grebes' greater diversity in bill form and Soot proportions which are associated with specializatiorls for taking a greater varicty of prey, may all be involved. Adaptations for pursuit diving, include larger size, which makes possible a relatively larger mass oSlcg muscles, lollgel-cnemial crests, which provide a larger area Ior the attachment of' these muscles, and the possible affect of the coiled barbulcs on the outer par1 of grebe's contour reathers, which act as capillaries in ahsorbing water, which rnay decrease buoyancy and may also cause this part of the combined featlicrs to act like a flexible skin which cause movement of the water in the feathers to act like tlie skin of a cetacean in produciilg laminar flow orwater across the surface of the birds. There is still much to be done before an adequate knowledge of' the m~~lticellrrlarparasites of' loons and grebes is known. The larger nuinbei- ofspecies of external parasites found on grebes (12 mites and 13 lice versus 1 mite and 2 lice on loons) is believed to have resulted Srom the association of grebes with coots and subsequent speciation on the larger number of species of grebes than loons. CONTENTS
PAGE
ABSTRACT ...... ii INTRODUCTION ...... 1 MA~ORSOUR(.F.S ...... 1 TIIEEVOI.UTION ANI) RELAXIONSIIIPS OF THE LOONS...... 1 TI~ENOMENCL~\.I.LIRE OF LOONS ...... 1 l'ul~~oses...... 2 Mr1.1rons ...... 2 TI-IE INTERNAL PARASITES (HEL>MINTMS)OF LOONS ...... 2 TI-IEI)I(.EN L.. ANS (FI.UIUS) ...... 2 THE(:ESTODES (I~\I~EWOI 1 Diversity of Loon and (;rebe Helminths ...... 22 2 Nurnhei-s of Nained Parasite Species Reported from each Loon Species ...... 22 3 Numbers and Percentages (in parentheses) of Hellninth Generalists. Specialists in other Groups. and Specialists in Loons and Grebes ...... 23 4 Loo11 and Grebe Melminths by Habitat of Iilfective Stage ...... 23 5 The Numbers of'Loo11Helrninth Species in Each Host-specificity Categoqr by Family ..... 24 G Major (;roups to which Final Intermediate Hosts of' L. oon and Grebe Helminths Belong .. 25 7 The Diversity of Helrninths in Loons. GI-ebcsand Alcids ...... 29 (Mourer-ChauvirC 1996) but is fragmentary and needs to be INTRODUCTION restudied; and Gauiella, the material of which is also fragmen- tary, whicll was tentatively placed in the loons by Brodkorb To datc, there has been no general list of the multicellular (1963), might be a plotopterid but requires further study (Olson parasites of loons comparable to that of grebes (Storcr 2000). 1985). In addition to the five living species, the genus Gavia is Becausc both arc foot-propelled diving birds that feed on aquatic represented by several fossil species dating back to the Lower animals, coinparisons between the parasites of the two groups, Miocene, where the early species Gauia egrriana of Czeclroslova- the spcciesof prcy taken, arid the biology of the birds arc of kia was found in the same deposit as Colymboides minutus (Svec considerable interest. This paper is intended to fill the gap and 1982). l'ollows the fcjrmat of the grebe work. It also i~rchtdesa conlpari- Long considered closely related to the grebes (Podicipedidae), son of the parasite faunas of the two groups and a discussion of the loons' similarities with birds of that group are generally the lictors that might have accounted for the diffcrenes in these considered the result of convergent evolution by Stolpc (1935) f>tun>ls.I liope this will inalie possible future comparisons with and subseqllcnt authors. This has been disputed by Cracraft the parasite laulias of other diving birds such as the alcids (1982), but see Boertnrann (1990) and Feduccia (1999:162). (I-Iohcrg 1984). Data from DNA-DNA hybridization (Sibley and Ahlquist 1990) Major sources. As with the grebes, major sources of informa- indicate a closer relationship with the Sphenisciformes and the tion on helminths are thc general studies of helminths, Procellariif'ormes, although loons lack the characteristic tubu- McDonald (1969); digciieans, Dubois (1968, 1970),Yamaguti lar ilostrils and strong, musky odor of the latter group. Small (1958, 1971, 1975); cestodcs, Dubinina (1966), I known from grebes. "Eupommtis " ~qbhosus,L. rnacrochiru.~"Heliopo-ca in,cisor;" P(~7-acoer~ogo,nirnu.sovat.us Katsurada, 1 9 14 I,. "Xc.notisn mgalotis, Micropterus dolomieu, Ponzoxis In Gavia .stellr~tn(J. Okulewicz 1984) Srnall intestine, "Huron sp.) , Cvuriiiidae (Notropis anogenu.s, N. FW. Eurasia. Intcrnied. hosts, see Storcr (2000). Para- atherinoidc~.r, N. cornuius, Pimephales "1lyborhyncliu.s" site of carnivores and birds of pl-cy, rare in other birds, notalu.~,Rhinichthys sp., Richardsonius sp., Semotilis including loons. [Recorded frorn Podic~f~scristntus in alromnculatu.~),Cvprinodon tidae (Fundulus dia/~h,anus), Ictaluridae (Amei,urus sp.), Percidae (l:'lheoslom,a HWML No. 38124). G. stellatn (McDonald 1969:84). "Boleo.soma" niyum, Per.cn sp., Stizostedion sp.) , Bursa Fabricii (most frequently, Gallimore 1964),intes- Sal~nonidac(Sa1rw.o sp.) (Yarnaguti 1958). Heron spc- tines. FW? Eurasia, N. Amer. Intermed. hosts? cialist (2), rare in loons, not known from grebes. Generalist (I), common in Lari, also in grebes, Alca, 'Iylodelf11~y.s g%os.soide.s (Dubois, 1928) Cepphus, etc. (McDonald 1969:84). Dubois (1968) di- In Gauin arclien. In ?intes~ine,?FLIT. Europe. "Colymb~~s vided Cotyl~~rusplatycephal~~s into two subspecies, "C. " a.vinticus" was listed in Yarnaguti (1971) as definitive I. communis, from Larus argen,tatus in N. Amer. and the host of Glo.s.sodif~lo,stomum(=Tylodrl$/~,ys) S;'ossoides. Be- nominate race from Eurasia. cause I cannot lincl "cr,sintic~i.s"inthe synonymy of spe- Order Opisthorchiforrnes cics names Sor either Gaviaor Podicef~sinthe twentieth Family Opisthorchidae ceritury. I think it presumably a 1af)s.u~calnmi for the Amphim,mu.s nrcticus Kontrimovitschus & Bakhmet'eva, loon, C. n.rcl%e;us. (Although, according LO Ogilvie-Grant 1960 1898, the specific name, "arcticu.~,"was sometimes ap- In G. stellatn (type host), G. immm (Kinsella & Forrester plied to the Horned Grcbc [Podicc$s n.uuil,us] in thc 1999 voucher HWML No 38108) and Conboy et al. in 1800s in Europe, whci-c the parasite was described. prep. voucher USNPC No. 087041.00), formerly listed Coly,mrOuswas the generic name used fhr the loons at as A. .speciosus is also this species vide Kinsella in litt.). the time $ossoid~ (21 01. 1985).Intestine. FW? Eurasia, N. Anlei-. Intermed. In G. steblata (vouchers USNPC. Nos. 073921.00 and hosts? L,oon specialist, rare in grebes (3).Formerly in 073922.00). Intestine. SW. Eurasia, N. Amer. Intermed. gcnera Armadoslr~jnbin,iaand Microsornacnnlkus. Taenia hosts? Paratcnic hosts? (;cneralist (I),also in Podice/x cnpilellaln Iiudolphi, 1810 (voucher in USNPC No. q7isegena, P cristntu.~,Mmpcs .reruntom; and Cepphu,s gry 11.. 035933.00 l'rom G. im,~ner)and Micro.somacan/h,us Formerly placecl in the genus Ilemiechin~osoma. Dc- psc,urLoroslellalu.s (Joycux & Baer, 1950) are synonyms scribed in Corynosomain which it is placed by McDonald ,/id(: Spassliaya 1966, and ,Joyeux & Baer consider (1969:661). I)~~cbl?~il~o/(~~is(/Mzc~~~om~~cr17cthus) sr~~io'er.sltii (Gasowska, AnrLracar~ll~aphalacroco7-acis (Yamaguti, 1939) 1932) a synonym of' ro.stellatu.s. In Cavia immer, G. srellata (Ryzhikov et al. 1985). Sniall D~ehin~i~n~ol@i,s.swirLe~slti% (Gasowska, 1932) intestine. Asia. SW. Intermed. hosts? Corinorant spe- In Ga71in nr(;liccc (?and pacqica), G. stc~llr~ta(Ryzhikov et cialist (2), also in Rissrr. Not known from grebes. De- nl. 1985). Intestine. FW? Eurasia. Intermed. hosts? scribed in Corynoso*na. considerctl a subspecies of ittrw~(~rinthe former USSR In Cnuin imtttm (Ryzl~iliov (el. 1985.) Intesti~le.FM'? (1)cmcnt'cv ot (el. 1968, original publication in Russian, SW? Fltrasia, N. iln~cr Ii~tc~mccl.hosts? Pnr,ltc~r~c 1950), the above rrcord of "imn~,c.r"almost certainly hosts? Also in grebes. Generalist (1) refers to od/rnt.tii. Similarly because (;nuicr /~nciJicawas C:o1-y17,o.rornnc.lnvtclrtrrr Goss, 1940 long considered a s~tbspecicsof (;. nrc.tic.c~,in both the In C~71instellnln (Rausch c:t 01. 1990). 111 i~ltesti~~e.FMI? hrnier USSR ant1 North America, the I-cport of (;. SW? Australia, Antarctica, Alaska. I~ltcr~ned.or "rrrcticn"co~tldapply to either species. If taken on the paratcnic hosts, fishes: Platvcc.phalidac (Platyc.r/~l~~~kts breeding grounds on the Arctic slope, it probably would Jitsc.trs). Spcciillist in co~.mor;lnts(2), I-are in loons, not be pnc.i/icn, otherwise it might be eithe~: (This ~mcler- known from grebes, also in seal (Gyp.so/~lzoc.n)and fox scores the value oSsaving specime~lsof'thc hosts, espe- (Alo/)pxlfegopt~s) (Klusch el nl. 1990). cially of' types.) /C;oryiro,soncrr .so~~t~trnc~(I'orssell, 1904) I'o1y1rrorphu.s ~ncrp~usSkrjahin, 1913 In (;/tui(r ittt,ttt(q;C;. s/ollntrr (Kyzhikov cjt (11. 1985). Intes- In C;uuin f/nlic.fr(and/or /)a(.j/ie(~),(:. .~l(~llnta(McDon;~ld tine, nrost oAcn at anterior part of large intestine. SW. 1969:67(i). Intestine. FMJ. l o11t that the finding of nicrnbcrs of this genus on plovers and Superfamily Analgoidea ovstercatchers (<:haradriidae arid Hacmatopodidae) supported Family Allopticlae my tentative conclusion (1956) that loons are related to BrPj,horc~/~sforJicigm (Megnin 8c Trouessart, 1884) ch;u-adriifol-rn birds. It has since hcen shown that loons are closer On G. i~nmm(T&e host), Gnvin nrcticn (Peterson 1971 ) . toproccllariifortn birds by Pragcr 8c Wilson (1980) and Sibley 8c A rarely found mite, know from a few collections from Ahlquist (1990) than to shorebirds, and Peterson's revision also northern Europe. shows t1i;tt species Ryf)horr~If~rarchund on many procellariiform birds. IJnfort~~niitcly,no phylogenetic analysis of Br~j~hosc~l~~rhas !rxt hccn madc, so wc cannot evaluate evidence of the relation- THE PHTHIRAPTERA (LICE) ships of thc loons froni the mites, although the presence of mites of this farnilv in Procellariifhrm birds mav be significant. Order Phthiraptera Order Aci~riformes Suborder Ischnocera Suborder Sarcoptiformeq Family Philopteridae 14 M~sc.PUBL. Mus. zoo^-., UNIV.MICH., No. 19 1 (~ras/~etlorr~rrtrusco1ymhirzu.c (Denny, 1842) crlburntr.s, Lruciscu.~lc~7cciscu.s, / FW. Annelids: LEECHES spp. Crustaceans: AMPI-III'ODS (;nui(r /)ac{ficcl Pacific I .eon Ganlrilariclac (C~arnmar.~c.ccf. locusla). Insects: DIPTERA Chironomidae (~hilan~omii.sislnndicus, 7h~nytal-,s~rsgt-ac.ilr?7~tu.s. FW. Crustaceans: ANOSTliACANS Family? "Aquatic insects" spp. Mollusks: GASTROPODS Lymnaeidae NOTOSTKACANS. Family? Insects: ODONATA Anisoptera (I,ymt7rrc,cc), Valvat idac (V(cl71ntn).Fishes: Cvprinidae (Alburn7r.s nyniphs. TRICHOPTEliA larvae. IHEMIPTERA Corixiclae spp. Mollusks: GASTliOPODS spp. Fishes: Gasterosteidae ((;c~~sl('rosl(?u,socrcl(:nlr~s, I-'un~gilizcs pzrngiri~is), Salmonidac (77~yrnallus Gnvln ndai~~s~zYellowbilledLoon ll!y?nnllu.s). SW. Crustaceans: AMPFIIPODS spp. Mollusks: FW.North (1994)reports the following food species available CEPHA1,OPODA 1,oliginidae (I,oligo o/~nle.scrns).Fishes: to this loon on fresh water lakes on the Colville River Delta, ~ttr;~clloidiclae(Poric1rlhy.s notntu.~),Clul~eidae (Cli~/jecr pollasi), Alaska: E~nbiotocidae(Cyrrmlogustc~ nggr(:gahs), Enoral~lidae(Engrnulus Crustaceans: spp. Mollusks: GASTROPODS spp. Insects: ,~tt,o'rdtrx),Stromateidae (Icicl~t1~y.sloc1~in~glorri.j. aquatic spp. Spiders: spp. Fishes: Cottidac (Myoxoc~j)halus Rcf'crcnces: Baltz and Morcjohn (1977),Davis (1972),Palrricr padricornis), Gasterosteidae (Pungitiu.~pungitius), Umbridae (1962). (Dallia peclornlis). SW. Annelids: POLYCHAETES, Nercidae (Nereis sp.) . Crusta- Gnvin int~rwConinlon Loon ceans: AMPHIPODS, Family? (0rchomonc:llasp.) , Family? (Anoi~yx n,irgnx), ISOPODS, Idotheidae (Irlothea sp.), DECAPODS, FW. Annelids: LEECHES, sp? Crustaceans: AMI'HIPODS, Hiuuolvtidae (Spirontocharis ochotensis), Payuridae (Pagurus sp.) , (;am~naridac (Gnvrnrnrr~slii~~nac:u.s), DECAPODS, Astacidac Pandalidae (Pan,dnlus danae), Mollusks: (sp.). Fishes: Cottidae (A.slncusk~itot~nc/ykc.s, Cntnbot-cissp.) , crabs sp? Insects: ODONATA, (12eplocotku.snvwzalus, Myoxoceplmlus joak?, M. scor)rrius), Gadidac Allisoptera nymphs. TRICHOPTERA, "caddis flies." HEMI- (Gadus morhua, iMicrogu(fus proxin~us).Cottam and Knappen PTEIW <:orixidae (Ar/:locorixrr.\tctilis). 1)TPTERA Cl~ironomidae (1939) Sound 11 5% of the stomach contents to be gravel. ((~lriro,rorrrrr.sislmndircls, 7i~ny/nmisgra(:%l(:ntz,sj.MoUwks: GASTRO- Refercnccs. Cramp ( 1977), North (1994),l'almer (1962). PODS l'lanorbidae (I'ln17~orOissp.). Fishes: Catosto~r~idae ((~nlnslmnrlscc~toslo~rrus, C. c.o,nnre~-so~ri),Centrarchidac (Amhlo/)lil(.r ru/ie,slri.c, Lel~omisgiIiDos~rs, L. ~~rclcrocl~i~ris,Micropt~rus .mlrnoidrs, RESULTS AND CONCLUSIONS I'o,rnox%s nrr~n~ulnris,P. ~n~igro~noculntus),Cll~neidac (Alosn fi.s~~~co'ol~(rr~~n~,r,rrcs,Ilmsomn ce/)~(iimn,z(:rr~),Cottidite ((;ott~tsDnirdi, In speculating on the possible causes of the differences be- L~$to/:olt~csarmatus), C;~vrinidae(C1~roso~nrc.s eos, HyDop~i~plz~mh~us, tween the parasite faunas of the loons and the grebes, I have Note,/togon,ls.schrysok(~uca,s, Notro]iis atl~r?.rinoides,N. /:orn~rkris,"Thick- included a wide range of possibilities, includi~lgthe ages, geo- head Minnow"= Isi~nc.l,hnl.:, promela,, Rhinichtl1,ys sf,., Scvrrotilir graphic distributions, and the degrees of morphological varia- c~tro.mc~/:~rl~t~rsj,(:vn1.i110dontiditc (fin(11tluxk~tero~lit~rs), Esocidae tion within the two families that might lead to different forag- (I:,sox lrcci~ts).Hiodon tidae (Hiorlon l~rgisus),Ictaluridae (Ameizcrzrs ing methods and hence to the taking of diff'erent kinds of prey n,(~Dulo.s~r,sj,1,otidac (I,otn lotnj, Pcrcidae (I'erccr flnue.ccen,s, containing different iinal hosts. S/izo.tleolioi~,ca17,adc~nse, S. vilreum), Pcrcousidae (l-'rrcop,si,s How good are the data? Several biases must be considered o~/ris/:ornaycu.s),Petrornyzontidae (I'elronlyzon rnarin,u.s), when analyzing the data. For the Common Loon, these have ?Plc~~ronectitIae"flo~~ndcrs," Salmonidae ((;orego,nris orlpdi, C. resl~ltcdfi-om differences ill focus of studies related to the food cbu/)e~[orrni.s,Saho grrirtln(~rii,S. trrtlla, Salvelinzts ,fonlinalis), habits, parasitology, and threatened status of the species in parts Un~bridae((lin1,r.c~ linri). Amphibians: URODELES "newts." of North America. Becarrsc loons feed prcdominantly on lishes, ANURANS "SI-ogs." both sportsmen and fishery operators often consider them seri- FW-SW. Fishes: Aneuillicbac (Anguilln an,guilln),Gasterosteidae ous competitors. Early food-habits studies focused on the sport (C(IS/(*)US~PILScrculc:n/,cr.s, l',r~rr.gitrcs ~rtng-iru.~) Osrneridae (O.sme~-UJ and food fishes and seldom included detailed identification of modox). invertebrntcs. For sirnibx reasons, studies on the pnrasites were =Annelids: PO1,YCHAETES Maldanidae sp. Mollusks: GAS- often focused on the role of loons as definitive hosts whosc in- TROPODS Mar~incllidae sp. PE1,ECYPODS sp. [ermcdiate hosts were these fishes. <:EPI4ALOPODS sp. Crustaceans: STOMATOPODS Souillidae 1,oons have great popular appeal, and the decline in the breed- (Sguilla sp.), DE(:APOI)S Cala~uidae(Cnlccf~po flamrnecc), iilg populations of the Conl~non1200n in parts of North America I-lomaridae (F1m11rnr1l.snmericn,nus), Leucosiidac ["Iliacanthidac"] has caused concern among ornithologists and conservationists. sp. Pcncitlae (P(~n/:~r.sdrcormntm), Portunidae (Callin,ectes,so/~idu~), In a study of die-off's of this species on the wintering grounds in Xan thidae (M(?iai/~/~emcrc.mcrrin).Fishes: Anlmodytidae (A,rnmodytc.s Florida waters, ICinsclla and Forrestcr (1999) found a greater cr~nc~rican~r~,.~),13;1crachoididae (Ol~scrnuspardu,.~), Clupeidae variety ol'specics and numbers of microphallid digenes whosc (Hrc.ooorlin ty,.rrn,n~rs,Clrtj)r?cr lraron,grts, Sjjmlkus .sfirattus),Cotticlac fii~alintermediate hosts are crustaceans in sick loons which they ((,'ol(rt.s.scor.f,ius, HerniI@idol~is h~r/1,ilel)idot11s, My~~xocr?./)hnl~u sc~rpixs), "thought to indicate a shirt in the loons' diet due to low fish E~nbiotocidac(Cyrnntop.sler crggregata), Gadidae (C;adzes callm-1a.h populations." They also pointed out the need for more studies G. ~~/ZO,~II~ILN,i\/leln~~,ogrammu.v c~egl.fin~is), Hacrnr~lidae (0rlhoj)risti.s on the loons' breeding grounds. At least in North America, the chry.so/)l(m),Merlucciidac (M~l(~n,g7c.smerlnn,ps.sj, Pleuroncctidae other species of loons breed at higher latitudes than most grebe (P/(~~~mno(.lr.s,/IP,YIL,s), Sciacniclae (Cynosciorr ,-(plus, Mzcrofiodo.nias species, and as a conseqlrcnce, their parasites have been less ~t,nrlulntus),Svnenathidae (sp.), Trinolidae (Eulnng%ngurnardus), tl~oronghlystudied. Zo:rrcidac (%orrrr.o,suivifinr.rr.s). There have been very few reports of parasites of grebes taken licli-rences: Bielsa and Forrcstcr, ~~np~~blishcddata, Univ. on salt waters. The major studies of grebe parasites in North Florida), CI-amp (1977),Fjeldsd (,in lilt.), Forbush (1925),Madsen America (cg.those oiGallirnore [I9641 and Stock [1985])have (1957),McIntyrc and Barr (1997),Palmer (1962). been on birds taken on the freshwater breeding grounds. TIIF. I',IOI.O(:Y 01;' LOONS IN l M.nlt~n . ' and Tale (1976). It might be added that the period. tarsometatarsus, the only k~~owilskeletal element of Neogneorni,~, At the timc cstimatcd herc for the separation of grcbcs Goln is onc of thc clc~nciltsmost likely to show convergence in diving their relatives, South Arnerica, Antarctica, and Australia were birds. Thc skull ("Polcrarnir"which has yet to be formally de- conilected as the continent, Gondwanaland. Up until the mid- scribed) reported by (:hatte jee (1997) to he a loon is definitely Mioccne, the climate of Antarctica was sufficiently warm to sup- not tllat ofa loon according to Feduccia (in lilt.), and after corn- port populations of grebes. So, in spite of the tenuous evidence, paring the figures in Chatterjce (1997) with skeletons ol'Recent I think it quite likely that the grebes arose either on Antarctica loons, I agree with Feduccia, but just what Polnrornlswas remains or South America, but probably not Aus~alia,because of its much to be clcter~nincd. srrra1le1-grebe fauna (three species, one of which, Podicef).~ Beca~~seall the other known reports of the Gaviidae are fro111 cristutus, was presumably a late arrival in Australia from the the Northern Hemisphere, the loons probably arose in that hemi- Palearctic via either Africa or southern Asia). sphei-e,and because to date, all the known records of the primi- Fresh or Salt Water origins? The closest relatives of the loons tive srtbfimily, the Colymboidinae, with the possible exception are considered to be the petrels (Procellariifhrmes) and the pen- of two partial bones from thc middle Miocene (Pungo River guins (Sphenisciformes). The considerable amount of morpho- Formation) of the l,cc Creek Mine of North Carolina (Olson el logical evidence for this is sum~narizedin Feduccia (1999 and crl. 2001), it is likely that the loo~lsarose in Eurasia. references therein) and is supported by molcclllar evidence Rccalrse Cooper & I'enny (1997) used Neogaeorn,is as the oilly (Prager &Wilson 1980, Sibley & Ahlquist 1990). Both the pet- report ofa bird ofa modern order to have existed in the Creta- rels and pcngt~insare excl~lsivelymarine, as presumably were ceous, its correct identilication to order is crucial to their pro- their common ancestors. According to Sibley and Ahlquist (op. posal that there was a mass survival of birds of rnodern orders cit.) the penguins split from the pcb-el-loonstock bcforc the loons across the Cretaceous-Tertiary Boundary. split from the petrel stock (by a difference in Delta T50H fig- The earliest known fossil grebes are those reported by Nessov ures of 0.4). Therefore, it is highly probably that the loons arose (1992) Go111 two Oligocerlc deposits ill Eurasia. Unfortunately, in a rnarinc cnvironmcnt. Modern loons still spend most of their this niatcvial has not bccn studied. The distal end of' a femur lives on marine or estuarine waters and undergo the molt and (i.onl the,John Day bcds in northern Oregon was described by regrowth of their flight feathers there. Shlrkldt (1915) as "Co~~mhus"(=l'odiceb.r) oligocarnt~s.Thcsc bcds The Red-throated Loon is a special case. Some populations arc now considered to be 01' lower Mioccne age (Olson 1985). breed on small moorland or montain ponds which are often Bccalrsc the gcncra of grcbcs were not worked out l~rltilmore too small and acidic to maintain sufficient food sources for the than 50 yeal-s arter this ft-agmcnt was described, its generic place- birds. Thus in Inany areas, Red-throated Loons continue to feed ment rcmains to be dctcrrnincd, b~rtit is most unlikely that it and obtain food for their young on coastal saline waters during will be i~lcllrdcdin I'odice/)s. the breeding season. This habit is made possible by low wing 011 the basis of tlle ~iumberofgellera and species and of grebes loading and structure of the Icgs, which are even more strongly now on each continent (Storer 2000), South America, with 5 adapted for pursuit diving than those of other loons (Boertmann qenel-a (2 endemic) and 9 specirs (6endemic), is the most likely 1990). These adaptations might be considered to represent a arca of origin olthe grebes. However, except for a qucstionablc stage in the cvolution of loons from petrel-like ancestors, but li)ssil rccord PI-om the 1,ate Mioccne of Florida for which no Boertmann presents evidence to show that they represent an details of the structure are given (Becker 1985), the most "primi- advanced coildition and that this species is the most speci;llizcd tivc" genus, Rollarr.dia, is confined to South hcrica. of the living loons. That Rollnn(lia t-ollcin,d is the species probably nearest to the In contrast to the loons, the grcbcs probably arose on bodies ancestral stock has been confirmed by a new cladistic analysis of fresh water. Grebes are unique in building floating nests usu- (FjcldsH ~ns).It and the genus Pol~ocephalusendemic to Australia ally anchored to the bottom, and this prevents their being moved and New Zcaland share similar courtship ceremoriies which, by winds or currents. The fact that all members of all genera of accortli~lgto detailed analyses, arc potential precursors of more grebes have this habit and rarely, if ever; build nests on land can elaborate behavior patterns of other grebes (Fjeldsb 1983,1985, be taken as evidence that the habit is an old one at least predat- ing the tlivision of the group illto genera. Brca~lsethese nests rrrinutu.c co111dstan(1 and eve11 run on land, it rniglit also forage are a11cliorec1to tlie bottom, they cannot be ~~sctlin tidal waters to sonlc degree on land. I doubt that the last was true. I li~iowof 01.other waters that lluctuate much in depth. For thcsc reasons, no rcpo~.tof any grebe actively fofirging on land and believe I believe that floating nests havc been a characteristic of grebes that all these birds (and (:olymDoid( Tahlc 1. Diver-sity ol Loon and GI-cbcHrl~l~inthsl Dige11c;tns Cestodcs Acan ths. Nematodes Totals All C;rcbcs 3 P~tlic.,,/).\spp.' All Loons GENERA All Grebcs 3 Podic.c.l,sspp.2 NI Loons SPECIES All Grebcs 3 ~Odi~($.\Spll. ' All 1,oons 'l~xlxcsscdin ~lu~nbersand (pcrccniagcs). I'i~diw/).ouri/rr.s, 1'. ,yrisc,g~?i(~,:~nd 1'. nigric-ollis (Mola~-c~icpopulaliorls only.) E~blc2. Nlul~l)c~.sof Na~rletlP;r~xsitc Species Reportctl from cacll ~ilajorgroups arr shown in Table 6. Many decapotls reach larger Loon Species' sizes than insects and small crustaceans like copepods and l>igcncans (:estodes Acanths. Nc~rlals.Ectopara. all amphipods and thus may be inorc sl~l?jcctto predation by looils (;crvin ,\lellrln 22 10 (i 10 1 49 than by grebes. The greater proportion of illsects as final hosts (;nvirr pnc.t/k(c 5 1 0 0 0 6 for grebes tha11loons is probably real in pu-t bccausc rnost grebes (;rruicr nlrlirr~ 8 (i 2 4 2 22 arc snlaller and because the Eared and Silvery grebes, Podice.l,s (;nuin nn.lic.cr antl/ol. nigricollis anti 1-1 occzl,ilalis, are less well adapted for pursuit div- pncl/ic.cc 6 8 4 4 2 24 (;nvicr irn~rrc,r 32 9 5 11 2 59 ing, but specially adaptctl for preying on small hosts. The greater (;nvin nd(o~r.\ii :'I 3 1 1 0 8 proportion of fishes in looils' tlicts is 131-obablya result of the loons' larger size ant1 preference for fccding on these animals. 'For a cornparable table of gl-cbc spccics, see Scorer (2000, p. 43). Ectoparasites. The numbers of genera and spccies of ectoparasites s11ch as mites and licc on ;I group of birds are in large part dcpcndent on the n~unbcrsof kinds of other birds takes advantage of the fact that the two groups have rol~ghly with which these birds have had physical contact that pcrrnits overlapping mngcs (Figure 2). (The parxsitcs of the Eared Grebe host switching to occur and on the radiation of the birds after populations outside of the Holarctic have not bccn st~~died.) receiving the parasites. It is also dependent on the hahitat in This reduction ofthe grebe parasite faluna to those ofthe three which the new hosts live and how this may afict the sltnrival of Holarctic species of Podic~psliasthe additional advantage of climi- the parasites. nating 11 species of ccliinostomatid digeneans and 6 of amabilid Twelve spccics of mites and thirteen of lice are known from cestodes kllow~ionly from the original dcsci-iptions from rrebes but only a siilglc mite and two lice Goiri loons. The dif- 7irchybc~I)tusr't1 ficollzs, Podilymbus /lodicc,ps, or Podic~;l,scrirtntus. fcrcncc in the nunlber of spccics of licc can be explained, at The relative diversity of' loon and grebe helminths is shown least in part, by the fact that ~lnrelatrdgenera of lice have on Table 1. The two fa~nilirs(the Amabiliidae) and tlie speciated on each gro~~p,Aqunnirn~us on the grcbcs and Dioecoccstidae), each with a single exception (not loons), be- Cra.spedonimus on the loons, and this has resulted in eleven spe- ing confined to grebes (Jones 1994) also increases the number cies on the former and two on tlie latter, a difference that can of genera confined to grebes by 7 and the number of spccics by be explained by the larger number of species and wider geo- 35. In contrast, but two genera (Biglnnclatriwrnand DuDi.n,inolepis) graphic range of grcbcs. asid no higher groups arc confined to loons, arid the former The greater ~lunihcrof lol~sespecies on grebcs is also ac- genus is vcry poorly known. counted for by grebcs' interactions with coots, which has resulted The n~unbersof'loo~l and grcbc parasites with final stages in in several species of lice s~uitching,more often, if not entirely, Gcsh-, brackish-, and salt-water habitats arc sho~vnin Table 4. from coots to grebes than vice vc.r~c~.For mites, the difference is The greater number and proportion of loon parasites with rna- less clear, but it might also be a niatter of past host switching rine life cycles again is, at least in part, an artibct of the relative from birds of other groups. The presence ofa hippoboscitl fly is amount or work done on birds kom the two habilat types. probably a result or the loons' nesting on land, which is ncccs- The relative numbers of Iinal hosts in animals of different sary for the life histo1-y of the fly to be completed, but tlie single Tthlc 3. The Numbers and Percc~ltages(in pal-entheses) of. record of this fly and the fact that it occurs on two fish-eating tlelnli~lthGeneralists (?I, I), Specialists in other groups, (?2, birds of' prey suggests that the record may be based on an iso- 2), and Specialists (?3-5) in Loons and GI-ebes lated occurrence. The association between the black fly, Simuliurn euryndminiculum, and the Common Loon, which is based a spe- Sl~ccilicitygronps ?1+1 ?2+2 ?3 thru 5 Totals cific attractant found on the loon (Lowther et nl. 1964), suggests that the association has been a long one. No comparable asso- Digene;rns ciation with these flies on other 10011s or on grebes has been Loons 35 (58) 11 (18) 14 (23) 60 reported, although I see no obvious reason why this associatiotl Grebes 43 (38) 26 (23) 44 (39) 113 in other loons might not exist if their breeding ranges occnrred within the range of' the fly. The greater nnrnbcr and variety of mites and lice on grebes also may be attributable to the relative ages of the groups, but perhaps more so to the lesser amonnt of physical contact be- tween loons and other species of birds which has presumably reduced the possibilities of host switching oP these ectoparasites. The greater amount oP time spent by loons on salt water, which is initnicable at least to mites, may also be a factor. The reason for the apparent scarcity of the feather mite, Brep/~oscebJi,r/icigthe only mite known from loons, is not clear, Nctn,~totlca but it may be related to two aspects of loons' biology. Mites can- Loons 12 (57) 8 (38) 1 (5) 2 1 not slin6ve i~nlnersionin salt water where loons winter; and loons GIcbc\ 1X (49) 10 (27) 9 (24) 37 molt their flight feathers simultaneously on these waters, the Red-throateds in the fall (Sept. to Dec.) and the Arctics in spring All GIolrps (Durinck et nl., 1994b), hence mites cannot move to adjacent Loon\ 73(58) 31(24) 23(18) 127(100) flight feathers as they do in birds that have a serial remigeal GI cbc\ 81 (33) (53(25) 105(42) 249(100) molt (Dubinin, 1951). From this it is probable that these mites Table 4. Loon ancl GI-cbcHclmirrtlis by Habitat oi Definitive Host Hcl~nintl~ 'Taxon EW' ?FW RM" SW1 ?SW FW& SW ?FW& SW Total All Grchcs Digcnc2uns 39 57 4 3 2 0 4 109 Cestodes 45 39 0 I 2 1 4 92 Acantllocepll. 6 0 0 2 0 2 2 12 Nematodes 15 17 0 0 0 3 1 36 All 4 GI-onps 106 113 4 6 4 6 11 250 % 87.6 4.0 6.8 Digcncalls 19 24 0 3 1 0 1 48 Cesrodrs 29 20 0 1 2 1 4 57 Acantlloceph. (i 0 0 1 0 1 2 10 Nematodes 9 14 0 0 0 3 1 27 All 4 GI-oups 63 58 0 5 3 5 8 142 % 83.5 5.6 9.2 All Loons Digencans 15 1(5 0 6 5 0 3 45 Ccstodes 10 8 0 0 2 0 2 22 Acanthoccph. I 0 0 3 2 3 5 14 Ncmatodcs 4 2 0 2 2 3 2 15 All 4 Groups 30 26 0 1 I 11 6 12 96 % 50.0 22.7 18.6 1 FW = fresh water, RW = hl-ackish water, SW = salt water. 2 l'o(li(,l,, cri~riluc,I! gri,\eg(!~rn,crnd i? nzgricollis. 24 MIX.PUBL. Mus. zoo^.., UNIV.MICI-I., NO. 191 Table 5. Tllc Numbers of Loon Helriiirith Species in cach Host-Specificity Catcgo~ylby Family Host S~ccificityCntcgoriea TotalSp.l? 1 2? 2 3? 3 4? 4 5? 5 (:athae~rrasiidae Psilostoniidae Philophthalmidi~e Echinostomidac Clillostornidae Schistosolnatidac Cyathocotylidae Diplosto~ni~lc Strigeidae Opisthorchiformcs Opisthorchidac Hetrrophyidae l'lagiorchiiformcs Renicolidae Microphallidac Prosthogonorrridae Eucotvlidae 1)igcnca Totals Total Sp.l? 1 2? 2 3? 3 4? 4 5? 5 Cestodes Psc~~dophyllidca Tctrabothriidca Cyclophyllidea Ces~odeTotals Acanthocephalans Polyrnorpliida Polymorphidae Nematoda Enoplida Dioctopliyrnatidae T~.ichliridac Strongylida Syrlgarnidae Ascaridida i\rrisakidae Spirul-ida Dracunculitlae Acuariidac Ancyracanthidae Dracunclidae Onchoccrcidac Neniatode Totals 'Categories: 1 = Generalisr. 2 = S ~ecialistin other groups, rare or occasiollal in loons. 3 = Specialist in loons, I-arc 01-unknown in other gl-oups. 4 = I Ex~xcsscclits Numbers: Digcncans Cestodes Acanths. Nematodes Totals Mnjor Grottp Grebes Grebes Loons Grebes Grebes Grebes Annelids 1 2 0 0 7 10 Mollusks 3 0 0 0 0 3 I)ccal~ods 2 0 2 2 0 3 Sm;tll Crust 3 1 .I 4 7 6 30 Insects 5 12 0 0 5 22 Fislres 15 6 2 1 8 30 Tcll.apods 3 0 1 0 0 3 Expressed ;ts Percentages: Digelleans Cestodes Acanths. Nematodes Totals Major GI-o~~p Grcbcs Loons Grebes Grebes Loons Grebes Annelids 2 0 0 7 4.5 10 Mollrlslis 0 0 0 0 9 3 I)cci~~x)cls 0 3 2 0 3 3 Small Crust. 13.5 6 7 6 29 29.5 I~lsects 12 0 0 5 3 22 I~ishcs 6 3 1 8 47 29.5 Tel lxpods 0 1.5 0 0 4.5 3 lTig111-c2. Map showirlg the World breedirrg distribt~tionsof the loons and of I'odicrps nurilus, P Lpisrgcna, and the Holarctic breeding distribution of I? .ni,gricollis, prepared byJohn Megahan from maps in Voous (1960) and Palmer (1962). (Breeding populations from Iceland, llic British Isles and the Magdalen Islands in the Gulf of St. 1,awrence are omitted.) either live in a placc snch as the dowl~ybases of featliers which Of the other 27, 22 are linown to be fishcs, two, snails, and one rrmains dry or tliat they move to s~rcha place before the n~olt each, oligocliactc worms, odonate nymplis, and anurans. takes place. Unf'ortunately, on what parts of which feathers these The parasites of thc five Palearctic species of grebes mites occrrr is not linown. Grebes also have a simr~ltaneousmolt (7hchybaf)lus r.~Qicollir\,Podicef~.~ auritus, P <@egena, l? crislnlus, and of the flight featliers, but in nlost species, this takes place while P n,igricollis)arc the best linown and the most frequently involved the birds arc on fresh waters. in the overlap. 7: ~~~licollis,P. auritus, and 1-I nic~'r.ollishave been The grebes' Iloating nests arc satl~ratcdwith water and ordi- Sound to carry 22,1-1 cri.status, 28, and P pis(?gf,~za,33 species also narily do not last fronl olle breeding se;tson to the next. Thus, found in loons. Although all take fishes, I? crislatus and some they do not provide a snitablc environment for parasites like populations of' P griseg~naspecialize in Gslies, 7: r~~lirollisis a ticks, fleas, bedbugs, and otliel- artl~ropodsthat havc stages in generalist, and I? a.uril,us takes more small invel-tebrates on the their liSe cycles reql~iringa pcriod on dry land. Loons often nest bl-ceding grounds and more fishes in winter. The smaller nnniber on dry land but keep adding wet niaterial to their nests through- fill- P rristalus than P. grisegerta may be accounted fol- hy the fact out the pcriod of' inc~tbation.Both loons and grebes are highly that the former is not founcl in the New World. On the snrface, aqmtic and havc little physical contact with birds of othcr groups. it may seem unlikely to find so many instances of grebes that a~-c The ~liiljorexception is that between grebes and coots (I'~rl%ca). not fish specialists being parasitized by lielmintl~swllosc final In their frequent and oCte11s~~ccessfill attempts to take over grebe stages are carried by lishcs. However; all grcbes take whatcve~. ncsts for resting places or as bases for their own nests, coots will prey is easily available (Storer, 2000: 45), and some fishcs may fight wit11 grebes, and rnixcd clutches of grebe and coot eggs act sll~ggislllywhen carrying parasites. A known instalice of this have bccn foltnd in soine nests. On the other hand, loons rarely is that of the stickleback, (;nsl~rosteu.c,which becomes slow-mov- nest neal; or have pl~ysicalcontact with, othcr birds that might ing when infected by the large larvae oi' Sel~i,stoc;(?,l~hal~~~solidu.~ pet-niit exchange of ectopa~asites. (Dolph Schluter, pel-s. comm.). This is but one of the many ways Single specics of two other genet-a of lice (I',\eudornenofio~zand in whicli, directly or indirectly, pal-asitesmake i~lt~r~nediatehosts I,n(?ttlobotlLn'on)are fc~undon grcbcs and have close relatives on containing then1 conspicnous or easy to captnrc for potential coots. Both genera contain several other specics on coots ant1 definitive hosts. otlier rallicls. This suggests that tl~elice on the grebcs were de- As pointed out above, loons and grcbes are not known to share rived hom coots rather than 71ic~umn. Two of tlic genera of any genera of external parasites. 1,oons arc not known to have niitcs (Rhinony.r.r,us and Neoboyn'aic~)found on grebes are also SI-equentphysical contactwith other kinds ofbirds or their nests found on coots indicating a similar sonrcc for these parasites on comparable to that between grebes and coots which is thought grebes. to havc resulted in exchanges of external parasites. The overlap in parasite faunas. Accepting the clrrrent belief Host specificity. (Table 5). Di~eneaus.Mclnhers of 4 orders, tliat loons and grcbes are not closely related (pp. I, 16), one 15 hmilies, 41 genera, and 60 species of digenetic trematodes can make several predictions about what helminth parasites they have been reported to parasitize loons. Of these, no genns is rnigllt share. First, that most of thesc parasites wo11ld be known only f'rom these bil-ds. Of the species folrnd in loons, 14 generalists in their definitive hosts, and, convcrscly, the ii-equency or 22 percent are co~lsideredloon specialists, and 8 of'these are with whicli they arc fou~idwonld be reduced in parasites that known only from the loon host. Loon specialists are known fi-om specialize on other groups of birds; second, that their final hosts all Sour of the ortlcrs and 7 (47 percent) of the 15 Families on would be Gequent prey to birtls of both groups; and third, that the list. there would be few, if any, parasites tliat would be specia 1'1sts on Menlbcrs of 3 orders, 4 Families, 18 genera, and 22 species of birds of both groups, or if this occurs, it would he the result of' cestodes have been reported to parasitize loons. Of thesc, 2 similarity of'diets. Tlie data bcar out these pr-edictions. genera (13ighnnclatri alld Uuhinino/(~)is)consist of loon special- Forty-eight species of Iicln~inths(22 tligeneans, 12 cestodes, li ists. Of the species found on loons, 7 01-32 percent are c:onsid- acantllocephala~~s,and 8 nematodes) have been reportcd fi-oni ered loon specialists, 1 of them known o~llySI-om the type host. both loolls and grebes. Of these, 34 arc generalists, 7 are spe- Loon specialists are known from I order and 2 fanlilies (the cialists ill otlier groups (3in anatids and 2 each in Ciconiif'or~iies Dilepididae and the 13ymenolepididae) on the list. and Lari), 4 arc grebe specialists parasitizing loons, 2 arc loon Ofthe 4 genera anti 13 species of acantliocepllalans known specialists parasitizing grebes. [One (Co1.1Jlvr~rincapillnns,) is said from loons, 1 species (l'olymorphu,~g(~71ii) is co~~siclcrecla loo11 to be connnon in hot11 grolrps, hut Utsileva ~l nl. (1999a) con- specialist. sider this species to bc a grebe sprcialist and that reports of this Mc~nbersof 4 orders, 9 families, 14 genera, and 21 species of species fi-on1other groups, incli~clingloons, to be "erroneous or nematodes have been reported Lrom loons. Of these, no genera doub6~11."] 1,oons are not known to be l~nrasi~izedlqr any of the and but one specics (Sjn,gumzrs arrticu.~)is considered a loon spe- (imembers of the Dioecoccstidac or 29 members of the cialist. Amabiliidae, which specialize on grebes. A single spccics of mite, the feather nritc (Brej)l~~l,hatceb/orJi(:%g.(tr), Fishes form the primary diet of'loons, that ol'grebes is more is fount1 on and confined to loons. It comprises one of six groups varied, althongh all species for which there arc sufficient tlata, of species in the gcnns (Peterson, 1971), but it is not clear to are known to take fishes, and some, like thc Great Crested Grebe which of the other five gronps it is most closely related. (I'odicej~srt-islat.us), the Great Grebe (Podicrj)horus mcq'orj, and the A single genus of lice (Crt~.rf)edon,irmus)is known from loons. It Western and Clark's grcbcs (Arch,rr~ol,lrorusoccidentc~lis and A. is confined to loons and is not closely related to otlier g?ncl-a of clnrltii) are fish specialists. The final hosts for 21 of the 48 spe- the Philopteridac. cies known to parasitize both loons and grebes are unknown. The black fly, Sin~uliumouryc~dmi17,iculzrm, feeds primarily on loons and is specially attracted to the Common Loon (Lowther and variety of potential intermediate hosts for helminth para- el ctl. 19(i4),although it occasionally will land on waterfowl (Fallis sites. Small prey are also taken in grcater numbers, which may, er crl. 1964).The hippoboscid, P,seudoYi.~-sin,furnip(!nnir,is not spe- in part, be responsible for the large number of individual cific to loons and may be only an incidental parasite on them. helminths found in grebes (up to more than 33,000) found in a lfwe accept tlie hypotheses that parasites which are generalists single Eared Grebe (Stock 1985), a species adapted for taking as rcgards their definitive hosts indicate a similarity in diet but small prey. not necessarily an evol~ttionaryrelationship and that the higher Boertmann (1990) has shown that the Red-throated Loon the t;lxonomic level ofa group of parasites specific to a group of diirers from all other species of living loons in having the lowest clelinitivc hosts, the longer the common evolutionary history wing-loaditlg both relatively and act~~ally,and most specialized shared by the parasite and host groups, we might be able to use legs for pursuit diving. The former is presumably an advantage ilil0rrnation on host specificity of two groups of hosts to esti- to these birds for moving from the small, oligotrophic ponds m;tte thc relative ages of thc host groups. where they nest to marine waters where they find the food Host switcl~ingSrorn one group to another may have occurred needed for themselves and their yoling. The latter presumably in tlic past, but it may be detected iS one species diSSers in its makes it possible for these small loons to compete s~lccessfully Ilost gro~~pProm that of the rest of the grotlp. For example, the with larger species and to take more pelagic than benthic fishes cestode Lliiiily Alriabiliiclar contains (i genera and 29 spccies than the Arctic and Pacitic loons. slwcific to grcbcs and one ~rionotypicgenus (Arnabilia) speciiic I11 underwater locomotion, there are the two variable forces to Ilaniingos, so it is probable that this Family's history was shared to be overcome: sizc, measured by the cross-sectional area, as with the grebes horn its beginning, and tllc species on flamin- mentioned earlier (p. 18), and the kind of flow (turbulent or gos rcsr~ltedfrom host switcliing. The case of the family laminar) of the water passing over the the body, measured by Dioecoccstidae is a little less clean In it there is but a single ge- the surFacc area of the bird. Because both arc areas, they vary in rills of 7 or 8 species, one of which is known only from glossy proportion to the square of linear measurements such as the ibises (l'l(~gnn'i.s) in Soutli America, whereas the other spccics, all length of the bird. On the other hand, the power, as represented of which arc grebe specialists, are known from all continents by the volume of the muscles used to move the bird, varies with rxcepthll-ica and Antarctica. This indicatcs that it is rnost likely the cube of linear measurements. Therefore, in birds of the same that this gelills (and family) originated with grebes. shape, the amount of power available will be relatively grcater Tlic paucity of genera and spccics of helminth parasites that in larger birds than the resistence or drag involved. Because arc sprcific to loons (2 geliera and 23 species) is in agreement turbulent flow caused by irregularities in the surface consider- with tlic relatively later appcarancc of the group compared with ably increases drag, adaptations that decrease this can have con- tliat of the grebes, in which the figures are two families or sub- siderable adaptive value in pursuit divers. In the case of dol- Fa~liilies,15 genera, ant1 105 sprcies (Storer 2000). The grcater phins and other cetaceans, turbulence is reduced in the skin by ncrmbc~-of species of parasites specific to grebes may also be in a layer of spongy tissue that holds water which can move as vari- p;trt 21 rcs~rltof the greater nlrmbcr of grebe species and the ations in prcsslrre differentially compress various parts of the Li~~riily'swider grographic tlistribution. The presence of two fami- skin's surface. Thus by making the surface of the skin smoother, lies or sr~blarnilicsspccific to grebes as opposed to none in the turbulent flow ofthe water passing over the skin can be damped loons is pcrhirps the greatest evidence frorrl the parasite fannas and result in a laminar flow (Hcrtel, 1969). In turn, this greatly oSthe great"' age of the grebes. decreases the amount of energy needed to move the animal through the water and permits the energy saved for use in in- WI IAT MI(;Il'l' Al:P'E(:'l' 'l'l-112. NIJMRER OF IIEL~lIN1'11SPECIES creasing its velocity. IN I,O~NSAND GREBES? A pec~~liarityof grebe feathers is the coiled barbules that lie parallel to the shaft of the barbs 011 the olrtcr third of the con- The size and speed of the bird. Perhaps the rnost sigriificant tour feathers. Although this was figured by Chandler in his dis- tlikrcnces ljetrucen loons and grebes with regard to their para- sertation on the structure of feathers (1916), he made no corn- site Saunas ;Ire the size of the birds and the relative nrlmbcrs of ment about the possible significance of these barbules. Fifty- two invcrtcbrates (V(>KSLLS fishes) in tlic diets. years later, Maclean (1968) described how the males of some Among predators with similar Seeding habits, the optimal prey sandgrouse (Pt~,ocles)use their belly feathers bearing similar sizc will vary with the sizc of the predator (Storer 1966). The coiled barbules, which act like capillaries, to carry water to their larger the prcy species, the fewer spccics of optimal size will be distant young. available. The sniallcr number oS prey species would mean the In moving rapidly under water, grebes hold their folded wings smaller number ofspccics tliat could act as final stages for para- against the body antl cover them with their flank feathers. In sites. 'Thcrcforc, the larger the predator the Sewer species of this position, the stiffwing feathers are covered by more flexible p;".;~sites it might be expected to harbor. However, this effect feathers containing coiled barbules on their outer third, and might be oflset by larger spccies of predators' being able to take other parts of the bird exposed to the water are covered with lax prcy of' wider ranges of sizc. feathers, the ollter third of which is similarly wetable. The barbs Grebes range in mass horn cn 100 to c.cr 1,600 g and loons on each vane of the feather and the coiled barbules along each from c.n l,(i00 to ccc 6,400 g with virtually no overlap between the side of the barbs lie at an angle to the feather and the vane, two groups. Tlie grcl~cs'smaller size makes it advantageous for respectively. Because this results in a "skin" of wet feathers with them to cake sriiallcr prcy, of which there are greater numbers coiled barbules lying in many directions within the plane of the and ~rricticsol'spccics. This in turn provides a grcatcr nrnnber feather, the "skin," by movement in the water in and out of the The smaller numbers and less variety of small invertebrates in Table 7. The Diversity of Helminths in Loons, Grcbes, and Alcids. oligotrophic lakes prcsu~nablymake feeding on these organisms Data from Hoberg (1984),Storer (2000),and this paper'. 2. less efficient so birds can better survive and raise their young there by feeding on larger prey such as fish which require Caster DIGENEANS p~~rsuitdiving to catch. Thus, it can be expected that large preda- Farrlilies Genera Spccics Loons 18 44 60 tors like Comnlon Loons use these lakes during the breeding Grebes 20 53 111 season, and that these birds will require large feeding territo- Alcids 13 22 29 rics because oS the sparser distribution ofthe available prey. This also nay have been related to the smaller clutch size in loons CESTODES (two) than in grebes (four or more). I.oons 4 16 22 As a result of these differences in habitat and the food re- Grebes (5 35 85 Alcids 5 11 2 1 sources in theni, birds nsing cl~trophiclakes will be subjected to a greatcr number and variety of intermediate hosts than birds NEMATODES on oligotrophic lakes. Loo11s 10 12 21 Birds spendi~lglong periods on highly saline lakes like Great Grebes 9 18 37 Salt and Mono lakes arc a special case. These lakes differ from Alcids 6 13 17 Sresh-water lakes in llaving a very small number of prey species, most ofie~ibrine shrimp (Arlemia),which rnay occur in vast num- A(-ANTHO<:EPHAL.ANS L.oons 1 4 13 bers. However, they are available only to birds like the Eared Grebes 1 5 13 antl Silvery grebes that are adapted to survive the concentra- Alcids 1 2 5 tions of salt (Mahoney & Jehl, 1985).Visits to Mono Lake, and probably Great Salt Lake, and swallowing the highly saline wa- ALL GROUPS ter, niay be advantageous to the birds in purging them of intes- Loons 33 76 116 tinal hel~ninthsthat may have been acquired on the breeding Grebcs 36 111 246 grot~nds.,Jehlhas examined many Eared Grebes taken on Mono Ncids 25 48 72 Lake and but rarely has found intestinal parasties in them. (Jehl, 1 Narried species only. 1988, and in litl.). On Icirgiz Lake in Ibzakhstan, there is a popu- Does not include two specics for- which grcbes act as intcrtnediate lation of the cestode, (h?z/luam'apodicipina, whose intermediate or paratenic hosts. hosts there are brine shrimp (Maimova, 1981). Because the parasite cycle is Ihund lhcre, the salinity of this lake is presum- life cycles are found in them are those with intermediate hosts ably less than that of Great Salt and Mono lakes, but I have been that move between fresh and marine waters. Most loons nest on ~rnableto find a figure for this (Storer 2000). oligotrophic fresh waters, spend more time in marine waters The changeover of parasites grebes acquire on the breeding than grebes, and are intermediate in at least the nurnber of grorrnds to those acquired on the marine wintering grounds digeneans. and vice vcma has been discussed in Storcr (2000) and presum- The Condition of the Definitive Host. An ailing bird that is ably also occnrs in loons, but proof that the degree of salinity not able to take its usual prey will take any other prey that is causes this remains to be demonstrated. easier to capture and thus inay take prey carrying infective stages Oceanic waters present difSerent situations for pursuit divers. of parasites that would be rarely, if ever, taken by healthy birds. I11 pelagic waters, the prey is often scarce over largc areas but A striking example of this is documented in the paper by Kinsella cl~unpcclin certain predictable sitlrations where schools offish and Forrester (1999) on the helminths of moribund or freshly prey on shoals of invertebrates and/or smaller fishes. The birds dead Common Loons found during die-offs on the wintering may stay near these areas 01; especially during the breeding sea- grounds in Florida. This paper is notable both for the large son, may have to return to them between visits to their nests or number of species of helminths found, especially for the largc yor~ng.IS the cotlccntrations of prey move, the returning birds numbers of microphallid digeneans for which crabs are the fi- niay find the prey by seeing the feeding activities of other birds nal intermediate hosts. This evidently resulted from loons hav- or largc fishes. Thcsc feeding flocks often consist of different ing to rely on crabs rather than faster-moving fishes for food. species of prctlatoi-swhich have different host-parasite cycles with Physical contact with birds of other groups. Because of their different components of the prey (Hobcrg, 1996). aquatic habits neither loons nor grebes have much contact with Other host-parasite systems can be expected to occllr in coastal other kinds of birds, and a result, they have far fewer kinds of waters and cst~~arieswith different types of bottoms. Such eco- ectoparasites than most birds. The major exception is that be- systems are pl-esi~mablymore stable but provide smaller num- tween grebes and coots, and, as mentioned earlier, this appar- bers of prey and thus are utilizable by individuals or small groups ently has resulted in host switching ofseveral kinds of mites and orpl-cdators. Because these ecosystems occur over long distances lice from coots to grebes. The possible sources of known species and often in isolated areas, I think it likely that relatively more of mites (1) and lice (2) on loons are unknown. of the pa~asitesin theni might be specific in their definitive hosts. The dil'Serences in the numbers of digeneans, cestodes, and PARASITE GV.NEKA I .OONS SI NRF WIT11 PETRELS AND PENGUINS nematodes found in loons, grebes, and alcids (Table 7) agrees well with the habitats occlrpied by the birds of these families. If the loons were derived from the petrel-penguin line, some Alcitls arc strictly niaririe and what helminths with fresh-water support might bc found in a comparison of their parasite fau- 30 MI.;^. PLIIIL.Mus. zoo^., UNIV.MICH., NO. I9 I ~ias.Five species ol hel~ni~lthsthat llave been l0111ltlill 10011s parasites from fi-eslily de;~clCommon 1,oons rlsnv11e1-e on tlie h;rvc also bccn forulcl in the petrels antl/or the penguins: the l~rccclinggro~uncls, where sampling of microbes ant1 examina- tligencans l)i/ilo.slo~~~uttrc~crilrnc~r~~~rr and Iel~~l~~~ocoi?l~~r~~.\ (miiczis tion for othel- possible causes of death too often have bee11 em- (Yam;rgnti, 1971),the cestodc Scl~isloc.(!/)I~~nlussolirlus (McDonald, phasized at the expense oS nletazoali pal-asitcs. (Mctliods Sor 19(i9),and the nematodes, Cycr/l~oslwt~~ccpI~~17i.sci and S/c~go/il~orirs collecting and preserving avia~ihelmintlis and protozoa can he tlio~~~etlocro(Ya~naguti, lS(i1). Tllc first thrcc arc gcsieralists in their Sormd in Dostcr and Goatcr [I9971 and rcfcrcnccs therein.) tlcli~iitivehosts, (:. phenisei is too little known to PI-ovidestl-ong I'arasitolo~~can be impor-cant in consel-vation. llowledge cvidcncc, ant1 S. cJitrrrreti~nt~,a pctrcl-pengrrin specialist, is rarely of what parasitcs arc found in a given spccics of Ilost, what tlic li)r~ntlill loo~isa~id alcids, whic.11 may he only incidental hosts. life cyclrs of tliese p;~rasitesare, and the parasite load a given 1,oons arc known to share at least 12 species of'helniinths with host can carry witlio~~tdamage to its well-bci~lgcan be crucial to alcids, as well as with birds of othel- groups, ~iliichis consistent saving a thl-eatcned 01-endangcrcd host species. Basic studies with the itlei1 that Sor pa~.asitcsof g~.cgariouslislles that attract that provide inSorniation oS this kiiltl sllo~rlclbe snadc while tlie mixed lieding flocks of birds, it wotrld be advancagcous to be host species have healthy pop~~lationsSI-om which a Sew indi- gc~lc~rlistsin their dcfiilitivc Ilosts. vitl~~alsc;un he sacrilicetl Sor this purpose. Only one species ofmite, the feather mite, Hr~~il~oscc~/(~,sJbr/i(igc?; For the grebes, I believe that the Sonthrrn Hemisplierc- lias lias bee11f'or111d 011 Ioosis. Mel~il)ersol'tlle ge1111s11;lve been l'o1111d the most to discover bec:ulse oSa probable <;o~~tlwanalandori- on 111i~lyproccllariiSorin and cllaraciriif'or11i birds as well as birtls gin of the group and because no Sossil grebes are as yet known ol'othe~.gro~rps, but as pointetl out earlier (11. Is),a phylogciictic fi-om Australasia and no fossils of the endemic spccics in So~~tli analysis of' the germs h;~snot been nl;rcIc so a close~lcssof' I-ela- America have been found. Althougli Ilhclz~~hc~/~tu,sc1onriniru.s and tionship betweell the species on loons and petrels cannot be l'otlilyrrrlius porlic~/).rarc known from the Pleistocene of Peru dcter~nined. (Carnpbcll 1943), 1 know of no repol-ts of carlicr grebe fossils At this point, 1 scc no strong cvitlcncc li.on~the pal-asitc litrl- f'~-omthat continent. nas to support a rclatio~lsllipof'tlic loons with any other grol~p The hrl~ninthparasites SI-on1the 6 endemic So~rtliAmerican ol'birds. gl-cbcs a]-calso little known, so far consisting only os4 digcnctic ti-ernatodes antl one acantliocephala11.The ~i~uiibersG-om tlie 3 endemic species of Arrstralian grebcs in.c not mr~chIligl~cr: 9 t~.csnatodes,2 cestodcs, ant1 5 nc~natodes.The paucity ofreports Thcrc is ~llr~chvaluable work is still to I)? (lone on the para ol'cestodes is particu1;rrly unfortunate in view ofthe large number sites of both the loons and the grebes. On tlrc loons, this ill- ol'specics of' this group Sound ill grebes ol' tl~eNorthern Hemi- cl~~dcss~u-vcys of hcllni~lths,especially on the ~narinewintering sphel-e (e.g., 11y Stock 1985), and the likelihood that the gl-ounds and the boreal breetling grountls irr Nol-tll Asnerica A~nahiliiciaeand tlie Dioccoccstidae may have ariscll in the ;uld on the lice cycles ofthe parasites. (Lif'e cycles are known for So~rthel-nHemisph?~-e with the grebes. sligl~tlyless thati one-liall' [ca. 48 per cent] of'tlic. hcl~nilitlispe- While we have a f%irlygootl ~lnde~.stantlingol' the rrlation- cics known rrosn the loo~lsas well as the grebes.) ships aniong the species and genera ot'grehes based on cladistic For parasites 01' both loons and grebes tlie ~najol-priorities analyses by Rochenski (1994) antl ~jelclsd(111s.) of mol-phologi- for Sirttu-c stndics in the Nortllern I-Iesnispheue are survry .cvork cal data, ;is well as niany beh;~vio~-alstl~dics by FjeldsB and 0th- to find out what parasites are ti)r~ntlin thcsc bir-ds ancl wol-king ers, Tve now ~~eedpal-allel molecr11a1-stl~dirs of I~otlithe grebes o11t of' lire cycles of these parasites. 111 doing this, spccirncrls of' ancl the parasites (cspccially tliosc which specialize on tliese hosts talien shoulcl also bc ~lscclto provide tissue saniples and birds). A comparative st~idyof the Miocene grebes, Miobnjilu.~ data 011 species of' prey taliell hy tlir hosts. In grebes the first TLI(L/~(T%and Thioi-nis ,sor.ic~~o,and the scattered Oligocc~lemate- priorities are finding o11t what spccics of parasites arc acquired rial repoi-teti by Nessov (1992) ~voultlI)e valuablr in placing the Li-om salt-bvatcr habitats and f'rom the less well strtdied spccics, early European genera within the fi-;lsncwoi-k of what we know the Pied-billed (Porlilymli~r.spodic~/is)and Least grebes (7i~c11~lic1fitus about the relationsliips ol'the Recent genera. 111 addition, the rlo~n%nicrls),in all habitats. FOI-the loons, the snajor priorities arc behavior of 7brl~yb(~/)tusrlominiciic, 7: ~roun(,hollr~nriine,7: j)rlz~lrri, studies of' tlie lied-throated, I'acific, 2nd Yello\v-billed loons on and Podice/1l~o1-7rsina~or rieeds much more study. ~rlariilchahicats and all IOrrr North American species 011 the 1 think the parasite gro~lpsthat show the greatest potential breeding grollnds. [I ~101rldemphatically not rccornlncnd cak- for Stit~trestudies arc the hiiabiliidae and the Dioecocesticlae ing Conrrmo~rLoons fbr this pllrpose o~itlie southern par-ts of with their nearly complete specificity lor their grebe hosts; and, the range of thc spccics where the birds are rare, but the total as Hoberg et nl. (1999) pointed out, this inl'ormatiorl cot~ld"be population is large enor~gliso that talii~iga sample where tlie ~lsetlfor estimating a minimum age for the radiation of the spccics is comlnon on the 1101-thernbreeding grol~ndsshol~ld cyclol~hyllide~~ns."Tllc largest gaps in 0111. knowledge of the do no Ilarnl, cspccially if~~oii-111-eeclersand failed breeders were Asnabiliidac arc the lack of material fi-om the South America sclectcd.] If'sl~chsindies were contlnctecl, the birds taker1 could and Australasia, to wllicli ~niostof'thc "pri~nitivc"genera ol'grebes ;~lsopl-ovidc tiss~rcsamples 1.01. DNA analyses, seasonal changes are confined, and information on the life-cycle of Anrn/iilicr ill pectoral ni~~scleand Sat mass relilted to niig~.ationancl the lnmc,lligPm.This spccics is confined to Ilamingos and s~tcllinlbl-- I~rccdingcycle, ancl study spcci~ne~lsf'or which there are few for matio~imight pr-ovide evidence as to whet11~1-the grebes or the detailed analyses of' geographic variation (Storel- 1988). Vallr- fl;nningos we]-c the 01-iginal hosts for the A~nabiliitlac. able data on food habits could also bc taken fi-on1the salrle speci- I nrgc that the A~nabiliidaebc the basis for a ~nrllti-pronged mens. Special cffo~.talso sho111d11c lnadc to collect mecazoan attack. First, the endemic South Americ;~n and Anstl-alasian grcbcs shoultl be cxarniilcd for thcse parasites and tlie life cp sized that the lriore one knows about the biology of a group of clcs ol'tllose f0111ld worked 0111. Then, a cladistic analysis oP the organisms, the sounder the rcsults oP cladistic and molecular ihabiliidae sho~~ldbe made and compared with that of the analyses, and even ecological studies, will be. There is no sltbsti- grebcs. SpeciIicity IOr species of grebes and geographic ranges tute for knowing whole organisms in the field and laboratory. of the l,al.asites sho~~ldbe determined. Finally, lacustrine fossil Documentation. In studies involving more than one grollp of localities li-on1 the larc Crclaccol~sthrough the Miocene (but organisms, documentation oP the materials used is particularly espccially the Palcocenc and Eocene) in southern Sol~th important. It is encouraging to find that voucher specimens of' America, A~l~arctica,and Australasia should be carcf~tllyexam- parasites used in recent studies arc being preserved in parasite ined for li)ssil grebes. Such fossils, if folmd, might providc cvi- collections and are available for the use off~~tureworkers. But it dencc 1.cgarding the proposed Gondwanaland origin for the is disappointing to find that comparable material of definitive grcbcs ant1 when this occurred. and intermediate hosts is not. It is especially unfortunate that For comparison with the Amabiliidac, similar studics of tllc doc~~mentationof the solu-ces of material for molecular studies Dioccoccstidae wolrlcl be ~~sefulin attempti~lg to date the origin is not provided whcn the identification of the source of this of thc 1;tttcr lamily. Althot~glitlie life-cycle on 110 species of material is cr~~rial.Even in s~rchwell studied groups as birds, I)iorc.oc.c~.v/~r,shas bee11 worked out, JOgis (1978) found evidence this needs to bc done. Taxonomic levels between species may that odol~atenytnl)hs are probably the internlecliate hosts for 1). change, as in the case of the loons parasitized by Po1ymor~l~u.s crs])~rinli.111-ope. Two good placcs in North hrncrica to worli out gnvi%(p. 10). Cryptic spccics arc being discovered, and consid- the lif'c cycles ofspccics in this genus would be so~~thcrnTexas, erable molecular differences are being found between wherc a liigh proportion of 'lirr1~yhr~filrc.rr1orninicu.s is known to bc populatio~lswithin species. Nor can M'C be confident that all ~~ara"tizctlby 1). rrcolj~lbrs,aud Alberta, where Stock found D.n,.s]~c.r voucher spccimcns arc corl-cctly identified, even in the best- in scvc~.alspccimcns of'l'oo'ic.c.l,., g~isegencr.(1985). The life cycle curatcd collections. Therefore, 1 urge that editors require docu- of I). /)(c)~icoifir0111 tllc glossy ibises in South America would be mentation of all the nlatcrial ~lsed. cspcci;~llyw~l~~;~l~lc to rompat-r with those of thc spccics of In these days, coauthored papers arc the rule, so I would go l)io~c.oc.en;t~t.sfi-o~ng1.cbes and might pl-ovidc evidence for whether even further in I-equiring tliat in thcsc papers, there be some his spccics or one SI.~IIIgrcbcs was basal to the origin of thc documentation ofwhich al~thorwas responsible lor which part(s) Lu~~ily.(The liltelihood that otlo~latenymphs are the intermedi- of the st~tdy.This is something the reader nccds and has the ate hosts of ~ncmbersof this fi~~nilysl~ggcsts the possibility that right to know in order to cvaluatc the work. Dio~c.oc.c,,shssplit off liom the i\nlabiliiclae.) Tllcn, nlolccular analyses of both the parasites and the birds "A little learning is a dangerous thing; sllor~ldbc made fol- comparison with those of the cladistic ones. Drink deep or taste not tllc Picrian spring.' In 1985, Stock inclutled a preliminary coevolutiona171analysis oI'the grel~esant1 the Anabiliidae in his thesis. The accumula- This advice is 110 less cogent now than it was when Pope tion of';lll thc ;111ove inPorm;~tion~vould rnalic possible a penned it nearly three hundred years ago. Students now call coevolutionary study similar to, bt~tevcn more complete, that lcal.11 the jargon oP cladistic analysis before they learn (iP ever) oP I-lobcl-g's on the Alciclae and Aknln~nic~(1985). how to write simple, direct, precise English. Similarly, thcy call Tlic tlivcrsity ill thc struct~ll-cofthe fvet in (living birds offers learn how to make cladistic analyses before thcy know the basic cxccllcl~tpossibilities for ftuictional anatomical studies, for in- biology of the organisms thcy arc analyzing. Cladistic analysis is stance, a comparison ol'tllosc of the less specialized grebes like an important rnetl~odolo~gyfor constructi~lghypotheses about 12ollrr~ttclir~and 7hrlt~~l)(z])ru,sspp. ~lit11the Great Crested Grebe or the pl~ylogen~ticrelationships within groups of organisms. It is the last with the cvcn morc specialized Western and Clark's based primarily on true principles that systematists have been grebes. S11c:h studies sl~ol~ldprovide evidence on how the grcbcs' using, consciously or unconsciol~sly,for a long time. These are loot strr~ctul-c(ancl tliat of Irl(!,rf)c.,nrni.cand its relatives) evolved that in cornparing the characters of diPPerent organisms one must and bccame divcl-silied, as wcll as how the patella became ill- polarize the states of each character (that is, tvhich is the more volvetl will1 thcse ~llccllanisms. "priinitivc" and which is the most advanced evolutionarily), and The decline in support for parasitology in this colrntry is dc- that cllaracters which are believed to have resulted from con- 13101-able,rspecially in 2111 age when biodiversity is being cmpha- vergent evolutio~~slio~~ld not be 11scd. From this, it should be sized. W1lc11 a spccics of ani~nal01- plant beco~nesextinct, all clear that the best analyses are those made by those biologists the spccics dependent on it ;~r-c: also lost. Yet too many admillis- who have the best basic knowlcdgc of the group being analyzed. tl.ators~jrrtlgcrcscarch more by the amount of overheat1 grants Not col-rcctly polarizing the states ofthe characters or rccogni~ I~ro~~gl~tto their i~istitutionsthan on the new ideas and infol-- irrg convcrgcnt evolution can lead to erroneous conclr~sions.If m;ltion tlic resulting research will provide. It is ol'course impor- the pi-emises are ~~ntenablc,cvc~l the best logic cannot prodl~ce ~iltli)r r.cscarcllcrs to raisc the f~rnds~leedcd. to co~ltlucttheir viable results. Similarly, because fossils are the only tangible evi- work, hrr~the amount raised shor~ldbc dctcrrrlined by the needs dence for checking the results of time of evoh~tionof organ- of the p~.c?jcctitself' and shoultl be ,juclged by the value of the isms, it is advisable that molec~~larbiologists engaged in this work tlati~:uitl itlcas I-cs~tltingTI-om it. should lino~venot~gli abo~~t the ostcology of the organisms they In cvol11tio1x11-ybiology, 2111 aspects oPthc biolo (;ingericll, 1'.1>. 15)SH: l';~lrol)iologic-a1~)crspccitvcs OII Mcsonyc.hia, Loon, (;civiri ~IIIII~~,on its rrintrring gro~urclsin Florit1a.J. Heln~intllol. Archacwccti, and tlre origi~lof' whale.\. (:llal)ter 15 in The EIII~P Soc. Wash. (i6(l): 1-6. gc~~ceof Whalcs,J.G.M. Thewisscn Ed., Plernum I'rcss, New York. ICi~lsky,El C. IWiO. 'The ye;~rlycycle of the Nort11c1-n Blue I'cngnin, (;owe(; W.(:. 19:1Y. A new ~enlatotleli.on1 the loo^^, Cnuin ~IIIIIL~,~;ant1 its l:'rir/yl~liilo~~iiiior ~~ouooholln~irlitrr/ in the \4'cllingcon Har-bout- at-e;~. rcl;ltionship to Hnc~iricilo/rc.l,l~u.\Jotli/~n, l.inton, 1928. Proc. LI. S. Natl. Dominion Mus. Rcc (New Zral;md) : 3: 145-2 18. Mns. 87, no. 3071, pp. 130-143. ICflie, M. 1986. Thr life-history of Mc,.\orc.lli\ d/,rilrc..ulali~t(Iludolphi, 1802) Elarland, W.B., R.L. i-\rrnstro~lg,A.V. (:ox, L.E. (;I-aig, /I.(;. Smith, Xc D. llict~,1909 (Trematotla, Echinostomatidac). Zcitsclin 1: 1';irasitcnk. G. Smith. 1080. A geologic time scale 1989. (:anil)ritlge Univ. I'ress. 7'2::3:35-343. (:ambridge, New York, Pol-t (:hestel; Melbo~u-ne,ant1 Sydney. I(ontri~novitschua,V. L., X '1 L. Baklrmrt'c\~a.11)CiO. [Helnlinths occur- Mcrtcl, H. 1969. I-lydrodynamics of Swimming and W;tve-ridi~lgUolplrins. ring in Guvio in the locvlantis of'tllr Iiiver I,e~la]T~.utly gclrnin~. Lal). Pp. 31-(53 iri The Bioloby of Marine Ma~nnlals,I I.?: Andel-sen cd. 10:12~1-18:'I.In Russian). iIc;ldenric l'rrss. New York ant1 1.ondon Icuroda, N. 1954. On the classificatio~r ant1 phylogr~iyof' tl~r01-drl- Hobcrg, K.1'. 1984. Systc~natics,~oogrogl.al,lly a~~tlrcology of platy- Tnbinares, particularly the- shrar~vaters(l'ii//iriioj, with special con- lleln~intll~x~lasites ol' the scabil-d lamily Alciclac (CIial.ad~-iifol-tr~cs: siderations on thcil- ostcology ;111d llilbit dilftl-c~~tiiltic)~~(Avcs). I-Icl-- Subortlcr iUcac). Ph. I). L)isscrrarion, liniv. M'ilsl~ington.324 pp. ald Co., Ltd., 'hkyo,,Japan. I-Iobcrg, E. 1'. 1987. llccognition of l;rrvae of the Tetl.at)o~l~riitl;~r(Euces- I.abcrgc, 11.1. A. &J.I). rtlcI.;~nghlin.1989. I<\~o//,llric/z//,c.ri (hphipocl;~) totla): inlplic;~lions1'01- the 01-igin of t;lpcwol-rns in mal-inc ;IS;in intcl-mediate host ofthc ncm;~todcSf~q/j/oc.rrrcr c.rrrc\iccr~ir/lo. <:an. homcothcrms. ibid. (i5:997-1000. J. 7,001. 67:23:15-2340. EIol~crg,E.P. ISS(i. F;IUII;IIdivc~sity among ;rvian parasite asse~nt)l;rges: I .;lmbrccht, I<. 1933. Handbllch tlcr l'a1aco1-nithologie.Berlin: (;ebrCidCr llrr intelxction of history, ccoloby, and 11iogcog1-;rphyin 1n;rrinc sys- Bor1111.;1egcn tems. Ur~ll.Scand. Soc. Parasitol. (i(2);65-89 I.eono\: V. A,, 0.I. Bclogurov, N. M. Sl~ag\.alcev;~,and S. K. 13onda1-cnko. tlobc~.g,E.I'., A. Jones, X R.A. BI.;I~.11949. Phylogcnctic an;llysis among 1965. [On ~hct~-c~natodr 1i1un;i ol.piscivorous ])it-dsin Kamtchatki. 1 111c I;~nrilicsof~hc (:yclophyllidca (Euccstoda) txrseti o11 co~rlpar;rtivc (In Russian.) pp. 130-158, plus ;rddendicln following p. 349. h Lconov, ~norplrolo~y,with 11ccv hypothcscs Ibl- co-c\wll~tionin vcrtch~-atcs.Sys- \I. A. [Papel-s on I~clnlintholo~y111-cscntcd to PI-of.A. A. Soholcv on tenraric Parasi~ol.4251 :73. the 40tll anniversary of his scientific 2111tlteaching activi~ics.](111 Rus- I-loff~nan,G.1,. 1967. l'arasitcs of Nortlr Anrcric.an Ircsl~w;~tc~-fishcs. Univ. sian.) Vladivostok. Not sern. (hlit Press, Berkeley ;tntl 1,os Angclcs. ix -1 486 pp. I.inton, 1.:. 1028. Notes OII tlic trcrnatodes of I~irds.I'roc. U. S. Na11. Mtts. I.Il~ntc~;W.S., & 141.13. Vernbcl-g. 1960. l'rcli~rrinary str~dieson the life 7:3: 1-:'di. histo~yof (,'codioc.rf)liolr(.,l)~~cr~id/.;iSziclat, 1928 (Tre~natotl~t:Stl-igeid;~~). Livezey, B. (:. 1988. Mo~-pl~on~ec~-icsof'lligh~lcssncss in tlic Alcitlac. Auk I. I'a~.asitol.4(<:797-799. 105:681-698. 1ntc1.11;ltionalC:om~nission on Zoological Nomrnt latru-e.1972. Extl-act I.owthe~;.J. K., X D. M. T4lootl. 1Wi.l. Specificity of ;I Black Fly, Siiritrlitr~ti from Opinion 981 in Br~ll.Zool. Non~cncl.20(1) 197'2: 15" 13ull. Brit. t~~tr~~a~/~~it~irrtcI~i~nl)avics, to\z~ardits host, tllc (;o~rl~no~~Loon. (:anatl. Or~lithol.(:lul, 92: 172. Ent. Yli:91 1-913. ,]c~I~I,,J.ll.,,]n1988. Biology of' the E;~t-cd(:rehe ant1 Wilson's Pllalaropc Madscn, b'.,J. 1957. 011the footl Irabits of some fi.\l~-rati~~gI)irds of Drn- in the no~lbrccdingseason: a study of atlaptations to stline I;rkcs. Stutl- n~al-k. Divel-s,XI-ebcs, mcl-ganscl-s, ;~nd ;luks. 1)anish licv. Game Biol. ies in Avia~~Biol. No. 12. iv t 74 1111. :3(2) : 19-83 Jiigis, V.A. 1978. [An unusual type of cestotlc tlcvclopmcnt in the Mahoncy, S. A., anrl,]. ll. Jchl, Jr: 1985. Avoid;rncc of salt-loadii~gby ;I dioccious spccics I)iorc.oc.c.c/u.\ o.s/~/.,-((:yclol~liyllitlc-;~: Acoleitl;ie) I (In diving bird 211 a I~ypersali~~rant1 alkali~~clakc: E;l~-cdc;~-cbc. Conclol- Esroni;in with English sun~mary.)Irv. ilkad, Nark Estonskoi (:(:K, sr~: 87:389-397. Biol. 27:31-37. Malakhova, It. 1'. l!)85. On the ccstodc f;ru~raof certain I;ur~ellirostl-;~l Jolransson, I..(;., Xc U.M.1,. Nol-l)c~-g.2000. Rsynrnrctric ~orsaitl IIII~~I-- birtls of IGrrclia. p. '32-104. h:Slrulil~a~~, S. S. , iund R. P. M;llakhowl watcr swilnn~ing.N;~tr~rc% 407:582-58:3. (Eds.) (in Ut~ssiiun).Ekologiyil I'ar;v/iticlicskikh organizunov Acatl. Johnson, C.W. 1922. Notes on distributio~lant1 11;lbits of some ot the Sci. k11-cli;lnHI-anch, I'ctl-o7;1votlsk. bird-flies, Hippoboscidae. Psyche F):7!)-85. Martin, 1..D. &,I. Tatc,Jn 1976. Tlre skclrton ol I~crl~/or~ii.\cld7~niu.s (Avcs: ,\olrnston,T.H. & L.M. Angel. 194 1.life Iiisiory oltl~e[I-cnlatotlc, Pf,trr.cig/,r Mcspc~.or~~itl~ifor~~~es.Sn~iths.<:ont~-ih. Palcobiol. No. 27: 35-66. nrulmlit 11. sp. Trans. Roy. Soc. S. Aust~:(i5:285-291. M;rximov:l, A. P. 1981. [Mot-phology and de\,clop~rre~~talcycle of the Joncs, A. 1994. Family Uioecocestitlac So~~thwcll,1930. I'p. :<<)I-398,and ccstotlc Coirjlunr-I// /~odici/~i~tn(Szyn~anski, 1905) ((:cstotla: the Family A~nabiliidaeBrnu~~, 1900. 1'1). 400-405, h ICli;rlil, L. F., A. Hyrncnolepitlitl;ie).] (111 Russian wit11 English sl~tntnary.) Joncs, and li. A. Bray. ctls. ICcys to the ccstodc pafitsites ofvrr-tct~~-;ttcs. Pa~.a~itologiya(Lcuingn) l5:325-331. CAli luteruatio~ral.Wallingli)~-d, Oxon. U.K. McDonaltl, M. E. 1969. <:at;ilogr~rof hel~llintlrsof w;~~e~.fowl(A~~;ltidac). Joyeus, (:11., Sc J. G. Rac1:1950. Sur quclqites espic.cs ~~ouvcllcson pert U.S. Dcpt. Intel-io~;Rlu-can Sport Fialicrics antl 14'Ildlifk.Spec. Sci. connucs tln genre Hyrnt,wol/.l,i.\ Wcinland, 1858. Bnll. Soc. Ucpt. - Wllcllifc 110. 126. M!ashington, D.C. viiit(i92 1)p. Ncrrclr;~tcloiscSci. Nat. 73 (sen 3) :51-70. McIntyrc,J.W. 1988. The (:o~n~nonLoon, spitr-it of'nol-thcrn I;~kes.Univ. IGnlcv, 1. 1985. On the rnol-phology, biolo North America, No. 121 (A. l'oole ancl F. Gill, Eds.). Philadelphia: Sinelail; N.R. l971b. Studies on the heterophyid tre~natodeApo/~hallus . . t\c;~tlc~~~yol Natural Sciences; M'ashingron, D.C.: The A~nerican01.- Irr~wis,tlre "sand-grain gru11" of' yellow pel-ch (PrrroJlauo~c-(~nJi.2. The ~~itlnologists'U~rion. metacerearia: position, strllcture, and composition of the cyst: lros~q; Otlcning, I<. 19(i3. Stl-igcid;~aus Viigcln dcs Bcrlinrr Tierpal-ks. Angew. gcogl-aphical distributio~xand variation. ibid. 50(5):577-584. p.,II'ISIL~I. .. .' 4(~3):17I-l82. Sinclair, N.K. 1971~.A reviewal of Odhnrrin odh11,n.lTravassos, 1921 Ogilvic-(;1.;1111,W.R. 1X!)8. <:;11;11ogucol'thc birds in the Kritisli Muscum, (TI-ematoda:Microphallidac).,~, Parasitol. 57:980-982. 26 (II;II-L)Order XVIII Pygopodcs. Pp 485.558. Skov, H.,,J. Durinck, M.F. Leopold, & M.1,. Tasker. 1995. Inportant bird Okulcwicz, A. 11-)89. [C~astr.ointcstin;IInematodes of Gnuin \lelln/a and arras Ihl-srabirds in the North Sea. BirdLifc International, Cambridge, (;rriiicr crrc.lircr (C;aviidac) 1 (in Polish with English su~n~nal-y.) . UI<. Wi;ltlomosci Pamzytologic~ne35(1): 35-42. Sltov, M.,& E. Prins. 2001. The i~npirctof' estuarine fi-onts on the disper- Olu~lcwic~,J. 1084. [Hirtl t~.clr~atotlesof Lower SIlesii~.11. Intestinal sal of piscivoro~tsbil-rls in the German Bight. Marirlr Ecol. PI-ogress trernatoclcs of Red-tliroatctl Divcl- (C;avi/r sfcllrcln (I'ont.). Wiatlomosci Ser. 2 14: 279-287. 1,.a~.~~ytologic~ne .. . 30(4): 503-510. Spasskaya, L,.P., & E.M. Ivakina. 1973. [(:estotlcs of Gaviifformcs, Olson, S.L. 11185. The fossil record ol'birtls. Pp. 79-238, F~I-nc~;D.S., Colyrnbifonnes, ancl Procellariifor~nesfrom Ko~yaksknational dis- J.R. 1 LIST OF LOONS AND THEIR MULTICELLUIAR PARASITES Parasites of Gavia stellata E~slron~gylidrstntuDijex (Nitzsch, 1819) Trichuridae DIGENEANS Barwcapilluria mqi (Madsen, 1945) Echinostornitlac Syngarnidae l CESTODES Diphylloho~hriidac 38 Mlsc. P~JI~I. Mus. ZOOL.,UNIV. MICI I., No. 19 I I)rg?a~n~ncriirl(,,rir,p/a (I<~rtlolplii,1810) Cotmocrphalus ohvrlnluc ((:I-eplin, 1825) I)i/)l~~~l/obollrriitrri~I~~IPIN~CIII ((:rrpli~l, 1825) Irrgirs~,ricrcirrollninn/cr (L,i~istow,1888) Tetrabothr-iidac Strepoccim cmssicarlda (Crcplin, 1829) fitrnbotlrriu~rrrcrerocc;l,hril~~r~s (Rudolplli, 1810) Dracunculidac I-Iymcnolcpididac Auiosc,r/)en~mocgooryr Snpr-ynga, 1965 I)ilbiniriol(~ism,\trlln/rrs (Abildgaal-d, 1790) Micro~on~ncnnllmpar-c~cornpre~,ssa (Czaplinski, 1956) Parasites of Gauia immer /Wicro.sorrrncrrriII~zis .S~~IIII~~S(JO~CUX & Racr, 11041) DI<;ENEIZNS I'silosto~nid;~c AC'ANTI-1OC:El'HALANS Psetrdof).\ilostomo utri-2unr (Linton, 1928) Polymorpliidac Catliae~nasiidac I'olly~norplrrts (~ccr~li.\Van Clcavc X Star~.ctt,1'340 Ribrrroin oitd~trr~~(P~icc, 1931) Poly))ro,;DItu. phrpps7, I(ost)~lev,1922 l'.chinostomid;~c N IMATODES I~el~zi~orlrritrn~rcsk?jnbrni 0slrln;u-in 1946 Dioctopliymatidac I(chi~iochrr.smrtsspinu1osu.s (Kudolphi, 1809) l:'rt,shur~~glidrst~tb;j?x (Nitrsch, 1819) I~cl~inrnlornarmolutr~rn (Fr-oclich, 1802) Tricliuricl;~c Hin~n,\thlcinlincia Dietz, 1900 Bo~u.sr~pillrtricirnc>r~i (Matlscn, 1945) ~LIrso)~.Ili\d~nficulal~~~ (Kudolphi, (1802) Anisakidac /\/licropary~hir~m/oce/um L)ie[l, 1909 Corrlnrrnecum r-rrdolpl~riHar~wich, 19(i4 Philophtlialmidae I'rirorhis actrnlh~~.~(Nicoll, 1!)0(i) (:linostomidae Clinoc/ornr~rncorn/)lancrlu~n (Ruclolplli, 1814) MITES Schistosomatidae Alloptidae A i~s/robill~c~nial(?rrigcilen,\irs Johnston, 11)17 Brc~~lto,scc.l~~,\,/or/icig~~~-(Megniri & T~rot~cssart,1884) UPndrz'tobill~ar*iapul7~m 7rl1~n/ci (B~auli, 1901 ) <:yathocotylidar Me,\ost~;l,hnnusa~~f~rr~dicr~ln/oid(~s (Price, 1934) Diploostomidae I)iplo.\/on~umgavium ((;ubr~-lct,1922) I'osthorLi~~loxlo~~~urr~~rrinimirm (Macalll~m, 1921) Paras~tcsof Gauia paci$cn and/or arctica 7jllotlrlpIty.\ ~mmer(Dubois 1961) Str-igeidae DIGENEANS Irhtlzj~oco/~lul-lorrrnficus (Rt~dolplii,1800) Echinostol~iid;~c Irhtl~j~ocot~~lttrusplalj~c~f)hnlus (Crepli~i, 1825) 1L.hinorl~ct.srnrlsrocrxc~lu.~ Dictj., 1909 Opisthorcliidae E(;hino/)arj~pl~irrrr~bcccirliis (Diesing, 1850) Anzphtnlc~~r,crr-cticr~.r lior~tri~~iovirscli~~s & Bakl~~rlc'tcva M(~.\on.hi.spo/yr~~.slrr.~(Dietz, 1909) Er.schouiort hu.\ linloni (Gowc~;1939) Microphallidae I'Iolniko71infodzrr1.\ (Linton, 1928) L(~)in.\wriol/rOrc~cl~~.vornc~ ((:replir~, 1837) Hcterophyic1;ic Eucotylidac /lpophallu, hrmjis Ra~rson~,1920 li~rrolyle?colrni Skrjabin, 1924 O'rjfitocotj~kroncciun (Crcplin, 1825) E,ircolIy(~ne/)l~ri/ic.n (Mclilis ill (:rcplirr, 1846) Gjf~locolj~l(~lin,qtn (CI-epliri,1825) I%rigic.olr longw. Ransor~i,1920 (:ESTODES Slictodorcr lnri/ormicolrr Sogandarcs-RrrnaI & Walton, 1965 Diphyllobothriidac Renicolidilc Scl~is/occ~hnlrr.ssolitlil~ (M~~cllo; 1776) Kc~r~icolnpollaris Ko~itrimovitscl~t~s& Baklin~ct'cva, 1960 Uilcpididac Microphallidac Lnl~r-ipoms,\krjnhini Matevossian, 1946 iMicrol,l~allr~~/orr~.\tm-iIGnsella & Deblock, 1997 N(?ovnli/)orczpnrnz.\/)lra (I.inton, 1927) M7crol,hallu\ nicolli (Cahlc & Hu~ininen,1988) Dact; 1944 Pclrcitlil(,pis itrcetrs (Wrdl, 1855) Otlhnmia odlrnc+ Travassos,lO21 I-Iy~~ie~iolepidi&e P1.ostliogonimidae l3iglnrzdntrr'trm blglcrndafriurn Spasskya, 1961 P7o,sll~ogonirn1tsovrcluc (Rudolphi, 1803) I)~rbi~~,i~~ol~;(,i.~/'ieI~r~r~nr~~~i(Sk1;j;lbin & Matcvossiari, 1942) Eucotylidae l)~rbi~rir~olr~~issroidr.r:skii (Gasowska, 1932) 'Ihnaitin /~dtsrllnrkoiSkjabin, 1921 iMierr~somci~crnllrlr~ pnr(urnirrosorr~ri (Gaso~vska, 1931 ) C:ESTODES ACdINTHOCEPI-IA1,ANS Dipliyllobothriidae Polymorpliidac Uipl~j~llohofhnumdi/rrmrcm (Crcplin, 1825) 1hIynior~~l~ii.sgcr71ii IU~okhlova,I965 S~lli~locef)l~a~~r~ssolidus (Murllcr; 1776) I'o!yrnorp/ln.s mrLgn,us Skt-jabill, 1913 Tetrabot111-iidae P~)OI?'INOI~IL~I.Vntinu~ir,~((;oezc, 1782) 7blmhoth1-ii~sr~rcrrr-orrf)holr(.( (Rudolplii, 1810) PoI~~~no~;D/~rr.sobti~sus Van Cleave, 1 91 8 Dilepididac O:yel~r,\tvrr~ibisop (Schll~idt& B~isIi,1972) NEMATODES Nmvalifjorc~porui.s{~inr (I.inton, 1927) TricIi1lridae Pcznr/(.rotc~rniarnrrsoini (Linton, 1927) IInr7isrrr/)illnrin ~nr6~1i.\(Rudolphi, 1819) Hy~ncnolcpididae Ac~~i~riidae Ur~$~nrrido/aeriinlr~icc!olatn (Bloch, 1782) Dtrbininol/,/ns psrrcdorosl~~/nL7~.s(Joyeux & Bae1;1950) IICE l)trbi~iiriol~~/~i,sros/c~lla/~ts (Abildgaard, 1790) Philopteridac Crcispedonir-mi~simmcr Emerson, 1955 ACANTI-IO(;EPElAI,ANS Polyrnorpliicl;~~ FLIES A~idrercrrntliagravida (Alegrct, 1941 ) Simuliidae A7rdmcnlitlia pl~alricrocornris (Ya~nagrrti,1939) Simuliurn c,uryadminir.ulu,m Davics, 1949 Cory~rosomcinn,ntrrrirtm Van Clcavc, 1945 Hippoboscidac I'olymor/~hrts/~rc?,i.\ (Van Cleave, 1916) PscudoIfe~sia/tlm$ennir (Sahlberg) Soritli~oc!l/inc~lri,spirla (Van Cleave, 1925) Parasites of Gavia adarnsii NEMATODES Dioctophy~n;llidac DIGENEANS EI/S~I-OIIU~~~~A/IL/)[/CX (Nit~scli, 1819) Diplosto~nidac Tric11uricI;tc L)zploslornum gauiztm (Gubcl-let, 1922) Rnr~r.srci/~ilke~rhmrtrgi (M;rdscn, 1945) Tylodelf~lzysimmer (Dubois, 1961 ) Syngamicli~c Strigcidae C,ynflro.\tornczplicnisci (Baudct, 1937) Iclt~lzyocotylz~rrrsWI-rrlicus (Kudolphi, 1809) A~risakidac (l~n/rccc.c~~ric~r,unrieger/l(l~~n (Rndolptli, 1809) CESTODES Aruariitl;~~ Diphy1lobothriid;le Cosrnorcphnlris ohvc,ln/t~t((:rcplin, 1825) L)i/~hyllobolhnu~nditwmum (Crcplin, 1825) I1/rmrntcrin ntlr17ir.n(<:rcplin, 1846) Tetrabothriidac Sl~~o/~Ii~or~r.\diom~tlec~e (Johnston & Mawson, 1942) Tetmbothrir~~rnarror~f~haluc (Rudolphi, 1810) S/r($~/oreir~~rrtrssica7ctlo (Crcplin, 1829) Hyrnenolcpididae S/rc;l,/orure MITES NEMATODES Alloptidnc Anisaliidae /~r~!p/i.o.$cr/e!.$/i~r~rlg(~r(Megnin & Tro~~essart,1884) Cwn/rncnc.cum rurlolphi I-Iartwich, 1964 APPENDM B AI)OENDA TO THF. PARASITES OF GREBES STORER (2000) New spccics dcscribcd. (Thcsc wcl-c not incl~~clcclill the n~nnbersof Vasilcva, G.P., B.B. Georgicv, & T. Genov. 1999. Palaearctic species of s1x:cies 11sctl in tl~c;trlalyscs in Storcr 2000 01-this paper). the gelltrs Confluarin Ablasov (Cestoda, Hymcnolepididae): rcdcscriptions of Conflziariu multisfi-into (Rudolphi, 1810) and Myn~cnolcpididae Conflucrriu,jc~po?iicn(Yamaguti, 1935), and a description of (~or~lunna (;on/l~inri(ckrc/b/~ei\iasileva, I(o~.nyushir~, ;111tl Gcnov. 2001h. In '/iccliybr~pt~~.\ sp. Systematic l'arasitol. 44: 87-103. rr~jirollir,type I~ost.This is the Conflunrinsp. in Vasilcva, Georgiev, and Vasilcva, G.P., B.B. Georgicv, & T. Genov. 2000. Palaearctic species of Gcnov, 1!)99. Srnall intestine. FW. Eurasia. Internled. hosts? the genus Confluaria Ablasov (Ccstoda, I-Iyme~~olepididac): ('o~i/lrtcrrinp,\~~~i~I~~fi~~-rifcrc~Vasilc~,Gcorgiev, and (;enov. 2000. In Podic$).\ rcdcscriptions of Conflucrricr podicipina (Szymanski, 1905) and (=7hclr~1hc~/1lzi~)rr~/irollis. Type host. 111 srnall intestine. El4'. Eurasia. Con/lnona .furrifem . (Krabbe, 18(59), and description of Conjluaria Intcrmcd. I~osts? /,tntd?furczb~nn. sp., a key and final comments. Systematic Parasitol. I)oll/ic.stl~!/~i.\gri.seg/:rric.lc\ Vasileva, I For ease in linding species ill the gcneral accounts of parasites and of loons in the the lists of prey species and parasites. pagc numbers for these are in italics . For the last two. the lowel- nu~nberis fol- the list of' prey species and the higher nrlrnber is for list of parasites of the loo~lspecies . Other entries al-e in plain text . Suhjcrts in the table of contents. the names of species of prcy and of interrncdiate hosts al-c not indexcd . The data on the last two are in analyza11le lorn1 on the web . (see p . 2) Acanthoccphala ...... 9 Rr($ho.srrb ...... 113 acantl~occphalans...... 9, 26 jorficiger ...... 2 13, 23. 26. 30 Acarifonnes ...... 13 brine shrimp ...... 29 acarina ...... 12 Acwwiidae...... 11 Acuariinac ...... 11 Cullinc!ctes .~@idu.\...... ii Adenophorea ...... 11 Carnallanina ...... 12 AeclrrnoB horus ...... 20 Capill/~narnergi ...... 11 clrukii ...... -6, 28 Capillariinac ...... I I occidnztali.~...... 26, 28 (:ardiocephaloid~sbra.nde.\ii ...... 5 Alcataenia ...... 31 ...... Cuirlior+halw ...... 5 Alcidac ...... 31 Oathaemasiidar ...... 3 ...... 13 Alloptidae ...... (brria1~eluelic.c~ ...... 4 Amabilia ...... 27 cestodes ...... 230 20 xnrabiliid cestodcs ...... cetaceans ...... 27 ...... Anlabiliid;~~ 19. 21, 22, 26. 27, 30, 31 chanqeover of parasites ...... 29 ...... Anrph~~nc:~*roarc.tic.7~~. 5 characte~.~,polarization of states ...... 31 ...... lintoni 5 Ciconiida ...... 19 sp(.c.ima\ ...... 5 Ciconiides ...... 19 ...... aniphipods ... 22 Ciconiif'ormcs ...... 126 anadro~nousfishes ...... 21 cladistic analysis ...... 31 Analgoidcw ...... 13 <.linostoinidae ...... 4 Ancy.acanthidae ...... 12 Clinostomoidea ...... 4 ...... il nrlr(~cnn.tlra graui(l(i 9 Clino.stomurri romplr~ncztvm...... 4 tnc?rgi...... 9 (,'olywiboi&.\ ...... l, 18 ...... /)halncrot.omci.\. 9 a7zqlic.u~...... I, 16 ...... Anisakidac 11 foraging on land ...... 18 Ano~~ro/nerriuciliata ...... 8 minvtr~s...... 118 tti icrclcrtn tlm...... 8 (;olymb.us...... 2 Aj)ophallus c~~neriramis...... (i "Colynzb~u"o1igoruc.nu.s ...... 17 b,n~i.s...... 6 (~on/lumincupillaric ...... 2 2 26, 39 doniczc.~...... 6 furcif~ra...... 39 rla~crrrsis...... 6 kr/~bh(~l'...... 39 nzurlilirigi...... 6 m~~ltz~tricrta...... 39 Aqii(urrirrnru...... 20. 22 podicipinn ...... 29, 39 ilrl~yLl~~r~orliyncI~~~s...... 11 pseudofuni/e(~...... 39 ...... i\nn.c~tlo.skjablnia 9 (,'onlrucnrc7~m...... I1 Ai./c:rr~ia...... 29 r~~dokhii...... II ;uthropods ...... 26 spiculigwr~m...... 11 Ascaridida ...... 11 .. aarieg~~lum...... 11 A~~stralasia...... 30 convergent evolution ...... 31 Atistralian grebes ...... 30 coots ...... 1 20, 26, 20 ilzistrobill~c~?zia ferrigalunsb ...... 4 copepods ...... 22 ...... i~oiosopmsgullinrdi 12 corixid bugs ...... 21 VLU.\~OVO~~...... 12 .. cormorants...... 19 Cmynoroma...... 9 anntarlum ...... 10 Daptornithidac ...... 17 cluvuti~n~...... 10 Uarriscccf,ilkuria rarf~oni~...... I1 ~(7rnrn...... 10 mergi ...... 11 .strl~mosum...... 10 bedbugs ...... 26 Co.smoc.~phalus...... 11 Biglcuridnlrium...... 9, 22. 26 diesin.@...... 11 bzghn~hlr~um...... 9 Jirlottei...... 11 Aillilcl~ziellapolonicn ...... 4 obr~~k~lllr~...... I1 hiodivel.sity studies ...... 32 (;otylurus aquauic ...... 5 Bittirim nlterrrcrlum...... ii platycef~liul~~~s...... 5 black flies ...... 4 23. 26 plalycephr~lus communi.\ ...... 5 Uluc crab...... ii c~abs...... 29 boobies ...... 19 Cr/~s~~edon.imas...... 14, 22, 26 rcj/y~r~.binu.s...... 14 immer...... 14 Cretaceous-Tertiary Boundary ...... 17, 32 Cryplorolylr co.n.cava...... 6 cqf)tocofyloi(Le.\...... 6 Echinochasmus ...... 3 lingua ...... 6 coaxulw ...... 3 Cyathocotylidae...... 4 skrjaDini...... 3 gcn.?, sp.?...... 4 spinulosus ...... 3 (:yr~ll~.o.\totnrrl)I~(~r~i.\ci ...... 11, 30 Echinof)aryf)hium6r~culu.s ...... 3 Cyclophyllidca ...... 8 Echino~tomcz...... 3 cyclophyllidcans, radiation of ...... 30 rmoluturn...... ? Cyrlzr.\ler(~ ilisue ...... 8 triuoluW ...... 3 Echinostomidac ...... 3 Echillostomiformes ...... 2 di~bchiclts...... 19 Echinostomoidea ...... 3 tlecapods ...... 22 ectoparasitcs ...... 29 Dc~n.drilobiharzirr...... 4 exchange of ...... 20 (tnczlin.rzrr~rn...... 4 Enulzornis ...... 1 pulvc~rctlr~~la...... 4 Enoplida ...... I I I)ir~.sirr...... 5, 6 Epherneroptera ...... 21 lligcnca ...... 2 Epistl~mium...... 3 digcn~uls...... 2629, 30 Ersschoviorchi~...... A, 6 I)ii:7"1""" ...... 7 8 linlmi...... 5,6 nl~or~trctu...... 7 E~~arnphimer~issibiric~%~ ...... 5 illl(.l'rtrj)lrr ...... 7 Eucestoda ...... 7 L)ilepididac ...... 8 26 E.~~cotylr col~ni ...... 7 dilcpidids ...... 21 nephritica ...... 7 Dioctophynlatidae ...... 11 Eucotylidac ...... 7 Dioctophymatoidca ...... 1I Eudyptula min.or...... 1 Dioecocestidae ...... 2 22, 26, 27, 30, 31 Eudy ks ...... 2 l)io~.c.~c;osl~i.~...... 3 1 Lupl(~0rni.s...... 1 flcolyll~Y...... 3 1 Gc.stron.gylide.\tu/)ij%x ...... 11 r~.sj)c.r...... 3 1 pr~r(crroi...... 3 1 Dipl~yllobothriidae...... 7 feather mite ...... 26. 30 diphyllobothriids ...... 21 -. reeding flocks ...... 29 L)iJjI~ylloboll~riumdihmcim ...... 8 Filarioidea ...... I2 Diplostomidac ...... 4 flamingos ...... 1.19. 27 Diploslorr~.u.m...... 4 fleas ...... 26 roly mhi ...... 4 flies ...... 14 ~/lf!xical~dum...... 4 fli~htfeathers. molt of ...... 23. 28 gcu~ircm...... 4 flow ...... 27 i.mrnc.r ...... 5 laminar ...... 27 m~crruycnse...... 4 turbulent ...... 27 j~.s(~udo.\/)alhacc.um...... 4 foot tracings ...... 19 sp ...... 4 hot-propelled diving birds. evolution of ...... 19 .sl,allrr~cc,r~rn...... 4, 30 Srigatebirds...... 19 Diptcra ...... 14 Fulica...... 126 20. 1)ivcr; I%larli-throated...... 1 Divcl; Grcat Northern...... 1 Divcr, Grccn-throated ...... 1 Diver, Red-tliroated ...... 1 Galapagos Islands ...... 32 Uivcl; Wliitc-billed ...... 1 Gasterosteidae...... 21 L)ocopl~omsnlricolor ...... 14 Ga~terosteu.~...... 26 bi.sci/o.s~rs...... 14 Gaviu...... 19 grc~srcrviccy).~...... 14 [ rrrctica] arctica...... 21 L)ollfI~.\ilr$~i.sgris(.gc.rt.icu~ ...... 39 [ r~rclira]paczJicu ...... 21 dolphins...... 18 27 1immrr] adanzsiz...... 21 Dri~cunc~~lidac...... 12 [ immrr] immw...... 21 Dracuncu1oitlc;r...... 12 ndumrii ...... 1, 15, 16, 39 drag ...... 27 arctrca ...... 1, 14, 37 I)r(~pcmirlol(c(~.r~iol(u~r~(~olala ...... 9 arctica uiridigularis ...... 1 I)ul)~n.bol.()i.s...... 22, 26 bor(,alis ...... 2 fI~hrrnrm.n.i...... 9 egmiana...... 1 j.s~~odora~lellat~~.\...... 9 zmmr ...... ii, 1, 13, 15, 16, 21, 38 immer elasson...... 2 l~llmttle...... 2 lril~poboscids...... 27 rn~~ollmri...... 2 I-lirudinca ...... I2 pnc.iJicc~...... 14. 37 llost switching ...... 2 22, 27, 29 pccc.i/ic.a and/or nrr.lie.n ...... 38 hosts. specificity categories ...... 2 phylogenetic alialysis...... 1 H~mrnolcpididac...... 9, 2 1. 26 sc?P 1c~nlrrorrnli~ ...... 2 hymenolepiditls ...... 21 .\lc,llrctr~...... l, 2, 14, 16. 21. 37 Ilymeno1ep.s ...... 9 stellr~tnsq~~cc~r~.uLr~ ...... 2 Hytlrirhic sp ...... I I superspecics of'...... 1. 21 G'rr7iielln...... 1 (>miidac...... 1 ibises. glossy ...... 27. 31 Gaviifornrcs ...... 1, 17 1thtl~yocotj~br:rt~sml~cus ...... 5. 30 Glossiphoniidae ...... 12 plulj~c.~fihal~i.\...... 5 (;lo.\.\orliplo~totr~.urn...... 5 Inglisc.ria rirmhamrrlrr ...... 12 Goirdwanal.und ...... 30 intrslinal hrlminths. number and varirty of'...... 21 Great Salt Laltr ...... 29 1schnocc1.a...... 13 (hebe, Clark's ...... 16, 26, 28, 31 Grcbe, Eared ...... 16 21. 22, 27, 29 Grcbc, Great ...... 26 Grebe, Grc;rt Crcstcd ...... 16, 10, 26. 31 Grebe, Hoary-hcaded ...... 20 Grebe, Horned ...... I 21. 28 Grebe, ZAcilst...... 2830 I.n~tnobothr.co17...... 026 G~.cbc,Little ...... 18 lakes (;rel)e, Pied-billcd ...... 28, 30 entrophic ...... 28 onllinc of foot ...... I9 oligotroplric ...... 29 Grebe, Red-nccltcd ...... 21 I.n/c~riporussk~jc~bini ...... 8 Grebe, Silveiy ...... 22, 29 I~rc.aiiiccpli.~lidac...... 7 Grebe, Western ...... 16, 26, 28, 31 1.ecanicephalidea ...... 7 grcbes ...... 1,26, 29 lerclirs ...... 12 bill f'orm...... 28 Lm~intmirllrchmchysoma ...... 7 coevolulionary analysis of ...... 31 lice ...... 20, 22, 29 DNA-DNA hyhridilati on...... 19 I.!gu/r~...... 8 lkathcl- eating ...... 20 eulywhi...... 8 Scatlrcrs, coiled harbnlcs in ...... 27 inleslii~alis...... 8 Tcct. evolution oS...... 20 monoLgrammn...... 8 flalrli Ikathcrs, molt ...... 20 Lig~ilitlar...... 8 floatiirg ncsts ...... 17, 20, 26 l~l)~dtoes ...... 19 fresll-water origin ...... 18 locolriotioii. undcrwatrr ...... 27 host specificity ...... 2 I Loo1i.A~.ctic ...... 16. 21. 27, 28 Miocene ...... 30 Loon, Commorr ...... 5 16, 21, 27, 28, 29 Oligoccnc ...... :30 (lie-olls ...... 29 ~'cllcts...... 20 Loon, Pacific ...... A,21, 27, 30 pyloric plug ...... 20 1.0011, Red-tlrroated ...... 6 17, 23, 27, 28, 30 toes, distal phalanges...... 20 I.oon, Yellow-billeti ...... 1621, 28. 30 toes, rotation of ...... 19 loo~is...... 2, 19 gruiforrn stock ...... 19 as interniediatc hosts ...... 2 gull-auk line ...... 18 hills of, Fig . 3 ...... 28 Gynrnophallidtc ...... 7 coevolntionary ailalysis of'...... 31 Co(vmbozdreli1te ...... 19 database. I~ttp.//~.mv.~~m~~~z.lsa.~~mich.eclu/curatoi~s/rws/...... 2 hahilats DNA-DNA liyhridiration studies ...... 16 grebes ...... 28 Sossil record ...... 1 loons ...... 28 jaw musculature ...... 28 Hc~~inalotr~phus...... 6 specificity catcgor.irs ...... 2 lodipns ...... 5 Harold W. Maliter I. aboraiory ...... 2 Hawaiian Islands ...... 32 M . pscudotemporalis ...... 28 If~rlpmicrl~ino.\omn...... 9 M(~rrlrtmrr...... 7 FIerni.slom.c~...... 4 sp ...... 7 f~~espcrorr~i\...... 1 19, 20, 3 1 sp., Lrolicrr group ...... 7 I-Iesperorniihiforn~cs...... 1 17. 19 mayflies...... 21 Hi~ti~rophyi!s ...... 6 M~lrmitta...... 1:) I-l~te~ophyidac...... 6 Mc.~orch.i\ ...... 3 Hc?tc~ro/)hyo$~.sis co.nlinl~cr ...... 6 dmtic.~rlr~tus...... 3 Flimns//~lanlinc.in ...... 3 polyc~,lus...... 3 Hippohoscidac ...... I< l~soselit~~~sr~ppendi(.~~Iatc~id~\...... 4 M(.lo,r.ltis ...... 5 petrels ...... 119 6ilis...... 5 Phaethontida ...... 19 irrl.rrrt~tliri.~...... 5 Pl~ngicolnlongus ...... 6 xonlhosomzts ...... 5 phalaropes ...... 120 Mic.i!o/)r~~j~~)liizirn/~L(.P~ILI~ ...... 3 Philophtlialrnidae ...... 4 Micropl-rallidae...... 7 Philoptcridac ...... 13 ~nicrophallids...... 29 Phthiraptera ...... 13 specificity ...... ii pinnipeds ...... 18 Microp11.1lloicIc~1...... 7 Plncobdrllu ornnla ...... 12 Micr~1~~1i.rrllrt.s...... 7 Plagiorchiata ...... 7 /i~rrr..~l(,ri...... 7 Plagiorchifor~nes...... 6 ~iicolli...... i 7 i'G;yac-lis ...... 27 rrir.olli,lire cycle, fig . covc~...... ii Plotnikooia ...... 5 s~J?...... 7 Jiodlens ...... 5 A4~c,u.~orncrc.n1i/I1I(.\...... 9 Plotornithidae ...... 18 /~r~rr~corrrl,r/,ss(~...... L) 1'odic.rpliorus m.njor...... 20, 26, 30 /)nrnmier-osnlnrr...... 9 Podire/).\ ...... 17, 20, 22 .smizll/lri.\...... 9 arcticus...... 2 Miobrr()h~s...... 20 mr~ril~fi...... 2 21, 26 rorrll~ri...... 30 cistatus...... l7, 22, 26 mites ...... 12, 20, 22, 29 yrir I'odiczpedic@t P(...... 20 I'orlilyrnhtu... fiodicef~s...... 22, 30 F)O~LTOT~I~J...... 17 Polioc~)hnlu.s...... 120 poliocrphnlzo ...... 20 Polymorphida ...... 9 Polporphidae...... 9 Po1ymorj1h.us ac1.iLgnri.msi.\...... 10 nculis ...... I0 I)os/./lcuLzs...... 10 In.mis ...... 10 gnuii...... l, 10, 26, 31 vlmagnzr.~...... I0 minut&\ ...... 10 obttuzo ...... 10 phi#,; ...... 10 Po~pocqllalz~ssp...... 7 Po.slhodl~)lo.\lomum miri.imiim ...... 4 prey size, optimal ...... 27 P~.orellariiformes...... 17 Prostho~onirnidae...... 7 u 1'rostho.g oronim~rso.ocr/z~.\...... 7 I'al.~c.tc.intliocc~~I~ali~...... 9 P,eudoltrtwoplr~y(~s...... G I'r~rnro~~rrogonir~~~isOUNIIIS ...... 4 h~udol/er~icl/utnipcnnis ...... 427 I'rircrc.rrcrricr crcll/.rrca...... I I Psm~dormopon...... 226 I'tri!rrdi/r~)i.s rtrc./.cr.s ...... 8 Pseudophyllidca ...... 'i ['(I i!(~/inlmbri(cria...... 3'3 I'.s~udol,silostom.u rjorium ...... 2 p"lXSitOlOh7 Psilostornidae ...... 2 dccli~icill suppor~of'...... 3 I Psilostornoidea ...... 2 inlporlance in conservation ...... 30 Psilostomurn...... 3 I'/lml/.lr.r~/~olI~.nn'l/..\lohrrlll..\ ...... 8 Pt/!ror.les...... 27 orirrr trtli.,...... 8 Ptiloxcnidac ...... 20 I'/ rricl~?i!olnenirrrcm.torrti...... 9 I'u/finu~ ~yiseu.\...... 18 I'(lr/~/!r/li.s ncnrilhiis ...... 4 pursuit divers, size ...... IS I-'rrr~~a/ri.~wtrsp?...... 7 Pygidiopsb qmalrc ...... 6 p.itclla ...... 19 summa ...... 6 Pcnguin. L.itlle Blue ...... 1 I~II~~I~IIs...... 7,19 . . pcnglllns, gri~rrt...... 18 pcntastonlids ...... 12 XeigIrllar(1ilin lomuiar...... 12 Ii/,l/l.c.g(~i!c.Orc~r1~r11.u~...... 3 slerr~cce...... 12 pctrcl-loon stock ...... 17 Rrnicol(~keimnhun ...... 6 ~x~~rcl-pcngr~i~~linc...... 29 pinyuis ...... 6 pctrcl-pcngrii~i-loonline ...... 18 pol1rzri.s ...... 7 Kenicolata ...... 6 Strongylida...... 11 Renicolidac ...... (5 Sulida ...... 19 resistence (drag) ...... 27 surface-volume ratio ...... 18 RI211~ony.~.v1~s...... 20, 26 Synganiidae...... 11 Klrynchobdellida...... 12 Syngamus arctictc~...... 126I, liibeiroia ondntmr ...... 3 llzomnsi ...... 3 Rollandia ...... 17. 20, 31 Z~cl~yyhr~p/~~\...... 18. 19. 20. 31 mllnnd ...... 7 20 dominicm...... 30. 31 rrovarl~ollandiar...... 30 pelzlni ...... 30 s;~lmonidfishcs ...... 18. 28 ruficollis...... 18. 2391. 22. 26. sandgrousc ...... 27 Tannisia frdtsclzcnI~oi...... 7 Sarcoptifornles ...... 13 Z~trin...... 21 Schi~tocrj~lz(~l~rzssolidl~c ...... 8, 26, 30 6irrmz.r ...... 21 Schistosoniatidae ...... 4 bi7~ntirrata...... 21 Schistoson~atoidea...... 4 Teleorchioidea ...... 7 Sciczdicnm rfigosn...... 12 Tctrabothr.idca ...... 8 scoters ...... 19 Tctl-aboth~iidac...... 8 Secel-rientea...... 11 tctrabothriids ...... 21 . Scuratiinac ...... 12 lc.lrnbol/m.u.s cf . lorulosus...... 8 . . Shearwatcr, Sooty ...... 18 zmmcr~nus...... 8 Sirnuliidac ...... 14 m/lf.ro(.e~)~l~l~us...... 8 Simulirim etiryarlmzn.ic1~171rn...... 13. 14, 2.1, 26 I~rrfidns...... 8 South America ...... 30 Tlzrromyzon " trizonard' ...... 12 South Atnerican grebes ...... 30 r.1 hiurnis ...... 20 Sorrthern Hemisphere ...... 30 socialn...... -0. 30 Soutl~wellinnIr~spidn ...... 10 ticks ...... 26 Sphenisciforrnes...... 1, 17 tonguc-worms ...... 12 Spil-nrida ...... 11 Trc~natoda...... 'L .Splrr~.didoJiluria/allis~~nsi.5...... 12 Trichinclloidea ...... 11 lik cycle ...... 13 Trichuridae ...... 11 Splendidofilarii~lac...... 12 tropicbirds ...... 19 Slcgophonzs diomechn(,...... 12, 30 7Iylodtlphys 7 ~lossozd~s...... 5 . C~ stcll~@olnris...... 12 imrwr...... 5 S/c!lln.ntrhasrn~~.sJalc(~lu.\...... 6 sp...... 5 .St($hunoprur(~...... 3 gifi~rli...... 3 p.set~doer.l~i~inta...... 3 United States National Parasite Collection ...... 2 spi7zosn ...... 3 1Jria ...... 18 sticklebacks ...... 21, 26 Slictod(, l(LriJormieol(L...... LJrinatur...... 2 .Sl,I;Dtocnrn...... 12 rrnssicaudu...... 12 rrassicntida cl~nmrlrii...... 12 Vnriokpic...... 9 ~ur~rroscw..sis...... 12 voucher specirnens ...... 31 prctinif~ra...... 12 rccla ...... 12 Slrigea nc(uoui.s...... 5 Wczrdium...... 9 sp...... 5 waterfowl ...... 18 Strigeirhe ...... 5 web-footed birds ...... 18 Strigeiformcs...... 4 whalcs, evolution of ...... 32 Strigcoidea ...... 4 wing-loading ...... 27 Swo1lgyl;lta...... 11 wing-propelled diving ...... 18