Bulletin WAX. HIST. SURVEY (^ tura.1 IIisto]:*3r Suii^vey The Crayfishes and Shrimps (Decapoda) of Illinois The Ubrary of ttit NOV 41985 Unive(»/,, u. It UrteiM-CiMinoaj^q Lawrence M. Page STATE OF ILLINOIS DEPARTMENT OF ENERGY AND NATURAL RESOURCES NATURAL HISTORY SURVEY DIVISION CHAMPAIGN. ILLINOIS VOLUME 33, ARTICLE 4 SEPTEMBER 1985 SIAIt OK II-UNOIS DtPARIMtMOF KNtROV ASH NA R RAL RtSOl R(.LS BOARD OK NA ri'RAl. RKSOIRCKS AND CONStRV A HON Don Ktciiison. Ph.D.. Chairman; H..S. (.iTnwsKV. Ph.D.. Snyrlajy. (:heitn\tt\: Robert U Me7< aij. Ph.D.. fiiohfisy: Wai TF.lt E. Hanson. M.S.. t.u^inreriti^: Lorin I. Nrvi.iN*.. Jr.. Ph.D.. Fotrstry: I,. 1.. .SrjOvi. Ph.D.. f>o/<»)fy. riiPoix»Rr- 1.. Broun. PhD . Hrprr- srulinii tfir i*re<,irrMr\ NATURAL HISTORY SURVEY DIVISION, Champaign, Illinois SC:iKN riKIC: AND rE<:HNK;AL S lAFK Paul (;. Risser. Ph.D.. CJnrf Amcf., K. Adam.s. Secretary lo thr Chirf SECTION OF AQUATIC BIOLOGY C.c)KI>^N. RoBKRi VV. Pn-D.. l(/NHf/( Rioloni^l nnd llrad Bi'CK, D- HoMKR. PiiD., .lijwilK liinlttiii\l I,ARiM(iR». R. Wfiiion. PhD. Aquain Riiilnii,i\( Sharks. RrcHARU K. Ph.D.. Aqunlii Rioloi>i\t Hf.hkndfjn. Robf.rt a,, Ph.D.. AssiHiatr Atftiatu Rntlo^isi pHH iPf. David P.. Ph.D.. Asfiotiate Aqualu fiiolni^ist Rfinboi D, Kfti'Rah a,. PhD . Awnnate Aqualu Binl<>si\t Ross. Phji iPCF., Pii.D . Aystxtatf Aqunlu /iioi(>iii\l Bavi.fv. Pftkr B., Ph.D.. Ay-^nlanl Aqualu liuilof'isl Hfnfbrv. Mtchaki. S., Ph.D.. Awistanl Aqunlu Builoi^i\l Hicks. Randai.i,. Ph.D.. As'.islani Aquniu Hinloiiist Horns, Wh.i.iam H.. Ph.D.. AsM\ianl Aqualu Bioloiiist I.l'BiNSKl, Kennfiii S., Ph.D.. Ay\islant Aqualu Binlo^isl Starnf.s, Wavnf. PhD-. A\si\laul Aqualu Biolotiist .Storc:k. IfdW,. Vu.U.. Awnlanl Aquafu Biolofiist Wn.KY, MicHAKr J.. Ph.D.. A--\i\tan( Aqualu Butlaf^ist San[)BFR<.er. Jf-Ns-Diftfr. M.S.. A\'-i\lanl Supporlur Si ifnli.\l Waitf. Jana I... M.S.. Assislnnl Supportn-r Si trnii.\l W'arrfn. (.arv 1. , B.S.. AsM.Uanl Supptntivf SnrtUnl Ai'sTFN. Doi'f.l.AN, M.S.. }u>u(n I'Tiilrwinnal SiifnliU Hoof,, M u:ha F-I. M.S.. Juuun l>Ti>frss,„nal St irnli.yl HiftFRT. Bfveri.f.y. Junior l*ii)fe\Mi)iial Surnlist KwAK. Thomas |.. M.S., Junmr Pmfrwinunl \iifttlt\t Pfrrv, l.ANCF-. M.S.. Juniur I'rnfr.ynitnal Si it'Ulisl SoBASKl. .StfPHES v.. B.S., luni'ir I'rulrwional Siirntist .Sons. tii<;FNK. Junior PrnfrsMotml S< irnlist Iazik. Pamf-1-A p.. M.S.. Junuir P^ofewional Siifnti\l Dovvi,iN(.. David, M.S., Tri imnal Assistant twiN(., Kathar^n I... B.S,, Tnhniinl Awistanl KoPHFiMAN, Jfffrf\. M.S.. Tfi liniinl Awistanl I.voN, Dai.f. M.S.. Tnhnunl Assislnnt Nf.wman. Barky t.. B.S., Tr-ihnual Assislnnl .ScHMiTTi.FH. Crak. D . M.S., Trifiuuat Assi\lant Wai.nfr. Ri'iH. Tr< hnnal A\si\lnnt {^I.Ai'S.SF.N. Ji'I.iF,. B.S., Junior Trihnual .-tssnlnnt C^RAWFORD, LoI'Ann, B.S,. Junior Tri linual A.wnlanl (;ros.sf.tt. Lorrik, MS., Junior Tfihnuat .-ixMslitnt DfQi'fnnf, Barbara, yuM/or Tnhnifal Assistant Das, Daviu, E.S.. Junior Tn hniial A\\i.\lnnt K1FID.S, Robert, H.S.. Juuioj Tnlinnnl A<r Trihnual Awistant MAtiNEl.lA. Stfve. B.S,. Junior Trihnual Awntanl Mai.mf.R, Sl'San. B.S.. Junior Trihnual Awislanl Mavfr. (jiRl.STiNF, B.S-, Junior Trt hnual AsM\tanl McDoWF.l.l.. Brian D, M.^.X.. Junior Trt hnual Assistant Mcl.CCKrF, AiAN D,, B.S, /i/n/ur Trt hint nl Assistant Mit.l.FR. Sli/.ANNF. M.. M.S.. Junior Trthnunt Assistant WiKF. I.VNN, M.S., Junior Tri hnii at .Assistant SECTION OF BOTANY AND PLANT PATHOLOGY (iRUNWAi.u, Ci.Ai'N. I'h.D., Botanist ami Hratt (iRANF.. J. [.ELAND. Ph.D,. M\i oloiilSl HiMFJ.lcK. K. B.. Ph.D., I'lanI I'alholoiiist Neei.y. Dan. Ph.D.. Plant I'nlholoi^ist .SciiOENEWF.iss, D, v., Ph.D.. l*liint I'alholngi.st Robertson. KENNnn R . Pn D , Botanist KNDRE.VS, Anton (i., Ph.D., Assoi tntr Botanist IvFRsoN, I.oris R., Ph.D.. AsMstanI Botanist Sfr(.fni. |a.mf:s K... Assoanlr Suppintwr Surnlist Nfiaon. Bfttv. Assistant Supportivr Surntist McKnk.hi. Bh I N., M.S.. Junior ProfrssKfnal Si irntist Bl'RtoN. Phhip, M.S.. Trihnual Assistant Harrison. Robert .*\., Trihnual .-tssisiant Hi'ANc 1.1-Shar. Ph.D., Tn hnual Assistant Karnf.s, Jean K,. M.S.. Tn hnual Assiytanl MoRAN. RoBBiN (:.. M.S.. Hrsrari h As.Kistant Noi.TE, David, B.S.. Hrsranh A.s.ti.Uani SECTION OF ECONOMIC ENTOMOLOGY Ri EsiNK. W'h I lAM (... Ph.D., Assoi intr t.ntoinohiiiisl and llrnil .\itifbn. Jami.s K , Ph.D., hntomolufiist .\KMBHtiM, Kdward J.. Pii.D., h.nlomotofiist Ko(.AN. MARt:os. Ph.D.. hnliniuilonist kt'Ht.MAN, DoNAio K.. Pii.I>.. hnlomoloiiist. h.xtrnsion Maduox. Josei'H v.. Ph.D.. l:ntotnoh}nnt Randei.i.. Rosr.oE. Ph.D.. UntmnototiiU. hMrnsmn B()ti>E\iAN, John K,. M.S., Assminlr hnlonioh^isl Kasiman. Catherine K.. PhD., .issm intr Kniomotonisl Kfi.soi. Alt.AN S.. Ph.D.. .4s\iHiittr hlntinnoloiiisl Irwin. Michafi. 1-... Ph.D.. Assimntr h.ntinnohmist Levinf, ti.i. Ph.D.. AssiMiatr hnlomologisl Stefffv. Kevin. Ph.D., .-tsso, B.S., Axsist. h\t. Sprc, Cornpulrr Applitalntns Ka.mpmfier, Gail. M.S.. Assistant Supparliif Siirrtltst l.EpAR. CfERAl D. Assistant Supfmrtn-r Surnlist McC.iFFFN. MiiTON K,. Jr.. ,\I.S., .4ssistant Supportn-r Stimtisl HaYER. Karen. .M.S.. Assistant Suppifrtwr Si irntist Roberts. Stephen J.. B.S., .issislant Supportn-r Scirntist .Swoeford. David. Ph.D., Assistant Suppojtn-r Siirnlisl Wai.sh, i.Al'RA. M-S., .-tssisiant Suppinln'r S* imtisl Da/ey. Doyi e. Junior Profrssumat Stirnlisl Post, SrsAN. Junior Suppurln'r Stirntisl (Carpenter. Sherla I.., \..\., Tnhnical Assistant i'.jssY. I.At'RiE, B.S., Trihnual .issistant PeteR-s. Brenda M.. Trihnual Assistant Sherman, Renee. B.S., Trihnual A ssisiani WiisMANN, Janet I... B.S.. Trihnual Assistant WiTiiROW. Jane. Trt hnual Assistant Morris. Marilyn. M.S.. yumor Trihnual A.s.sisiant Mc(;iiRE. .Michael. M.S.. Junior Trrhnual Axststani CtRlDER. DntolliCii. Tran.si-ribinfi Srtrrtary, l:\lrnMtttl Lrdman, Diane. (.Irrk-Typist III 1 RAiB, Joan. (Irrk-T\pist III S/KonxiNsKl. .\\.\c.f,(.lrrk-Typisl II BaI'CHMAn. I'homas \.. (iraduatr Hrsranh As.sislanl .\rif, Syed Moha.mmad, M.S.. (iraduatr Hrsrarch A\st\tanl Day. tRic. M.S., (iraduatr Hrsrarch .Issisiant SciiRoEDER. .\i.AN, M.S.. (iraduair Hrsranh .^vw.w«n/ Smith, I.ane. M.S.. (iraduatr Hrsrarfh .-txiislant KiELDiNc, Dennis J., (iraduatr Trathin^ Assistant SECTION OF FAUNISTIC SURVEYS AND INSECT IDENTIFICATION LaBergf. Wali A( e t.. Ph D,. Insri-t Taxonomisl and Hrad Brigham. Warren l\, Ph.D . Insrft Taxonomist Pa(.f. Iamrencf M.. Ph D . Fish Taxonomisl Webb. Donald W,. Ph.D . Insnl Taxonomisl Bric.ham, Allison R., Pii.D . .issonatr Piofrssional Stirnti.st Godfrey, (>eor(.f I... Pil D.. .4\si>tiatr Insrtt Taxonomisl I'nzicker. John D , Ph.D.. .isAiu lalr In.snt Taxonomisl Von;riiN. David J.. Ph.D.. Assi.stant Insnt Taxotutmist rREWoRf.i. C^oLlN (;.. B.S.. As\4uialr SuppttrtuT Sitrnlist Bi'TCHER. Matthew K. M.S., A\Mslant Supportnr Sarnlifi Gardner. Genf. M.S.. .issi.stanl Supporln-r Stirntut HoEMANN. lovcE, Ph.D.. Assistant Supportit^ Siirrtltst McfiiFFEN. Katiirvn. M.S.. Assistant Supporln^ Sumlisi PFRAIT. Si'ZANNF J., .issistant Supp/M»rlnr Sarntist Kasprowu/. jE\NiNf M., B.S.. Junuir Suppo^tnr Stirntisi Kefnf, Dennis. B.S.. funun Supportnr Surntnn Mai mboR(., Pmti I.., MS., Srnior Trt hnual .i&sislani Bkowir. Joan 1... B.S.. Tn hnual Assistant HtMH)N. Steven. ,\I.S.. Trihnual Assistant Kffvin, Khomas M.. M-S.. Trthnuat .i\si\tant KiRTN, .\nn, B.S. Trihnual .issistant MiiF.s, Chari.ene. Trihnual Assistant PosMiA, SiE^F. Trt hnual .issistant Sw ANSON. C^iiARLE.s. S.. B.S,, Tri hnuat Assistant \ U.S LSSN 0073—1918 Nra.tii.ra.1 History Sixrvey The Crayfishes and Shrimp (Decapoda) of Illinois Lawrence M. Page STATE OF ILLINOIS DEPARTMENT OF ENERGY AND NATURAL RESOURCES NATURAL HISTORY SURVEY DIVISION CHAMPAIGN, ILLINOIS VOLUME 33, ARTICLE 4 SEPTEMBER 1985 Frontispiece: Orconectes illinoiensis form I male from Gibbons Creek. Herod, Pope County. Illinois. 4 March 1973. i ^nis' tepo^i/ i&' dedicated to/ MRS. BERNICE SWEENEY o*v tke^ occasion' ol k&i/ leiUement/ and uv konon/ ot ne^ 22 uewi& at se^dce^ toy the/ Section/ ot'^aunisUo SiMueus' and/ 'insect/ 'ddenliiication/ ol tAc/ ^(jtinois/ '^yialwiai ^~}(AstjOAij/ SiAAMeu/. '~^teA/ numetous' ketplut cantuAutions' to/ this/ and oth&i/ studies' OAe/ sinceAeiu/ ofipneciaied. Ill CONTENTS INTRODUCTION 335 Historical Studies of Illinois Decapods 335 Present Survey Methods 336 Extralimital Species and Studies 336 GEOLOGICAL HISTORY OF ILLINOIS AND ZOOGEOGRAPHY OF NATIVE ILLINOIS DECAPODS 339 NATURAL HISTORY 343 Crayfish Life Cycle 343 Freshwater Shrimp Life Cycle 345 KEYS TO ILLINOIS SPECIES 345 Key to Decapods of Illinois 348 Key to Illinois Crayfishes, Based on Form I Males 353 SYSTEMATIC ACCOUNTS 356 Family Palaemonidae—Freshwater Shrimps 356 Genus Macrobrachium Bate 356 M. ohione (Smith) 356 Genus Palaemonetes Heller 358 P. kadiakensis Rathbun 359 Family Cambaridae—Crayfishes 362 Genus Cambarellus Ortmann 363 C. shufeldtii (Faxon) 363 C. puer Hobbs 366 Genus Procambarus Ortmann 369 P. gracilis (Bundy) 370 P. acutus (Girard) 376 P. viaeviridis (Faxon) 378 P. clarkii (Girard) 381 Genus Orconectes Cope 386 O. illinoietisis Brown 386 O. immunis (Hagen) 388 O. indianensis (Hay) 394 O. kentuckietisis Rhoades 396 O. lancifer (Hagen) 400 O. placidus (Hagen) 404 O. propinquus (Girard) 406 O. rusticus (Girard) 412 O. slannardi Page 415 O. virilis (Hagen) 4 1 7 Genus Fallicambarus Hobbs 422 F. jodiens (Cottle) 422 Genus Cambarus Erichson 426 C. tenebrosus Hay 428 C. rusliciformis Rhoades 432 C. diogenes Girard 434 C. robustus Girard 439 CONSERVATION 440 ACKNOWLEDGMENTS 440 LITERATURE CITED 441 INDEX 447 Page, Lawrence M. 1985. The Crayfishes and Shrimps (Decapoda) of Ilhnois. Illinois Natural History Survey Bulletin, Vol. 33, Art. 4. p. 335-448. The Crayfishes and Shrimps (Decapoda) of Illinois Lawrence M. Page Crayfishes and shrimps (Decapoda) are conspicuous components of the aquatic environments of Ilhnois. Al- though only 23 species are known to inhabit the state (Table 1), they are among our largest aquatic invertebrates and often are present in large popu- lations. HISTORICAL STUDIES OF ILLINOIS DECAPODS Most of the historical information available on crayfishes and shrimps of^ Illinois is that published by Forbes (1876), Rietz (1912), and Brown (1955). The "List of Illinois Crustacea" was, as Forbes (1876) noted, "a first contri- bution to the knowledge of our Crusta- cea," and contained the "results of a single season's work." Included in the list were Macrobrachiurn ohione, Pal- aemo7ietes kadiakensis (as P. exiUpes), Procambarus acutus (including Cam- barus stygiiis), P. gracilis, Orconectes immunis, O. propinquus, O. rusticus (as O. placidus), O. virilis (including O. wisconsinensis), and Cambarus diogenes (as C. obesus). Hagen, in his earlier (1870) monograph on North American crayfishes, had noted the occurrence in Illinois of all of the above crayfishes except P. gracilis. Hagen erroneously also recorded for Illinois P. troglodytes, an Atlantic Coast species. A bachelor's thesis submitted to the University of Illinois in 1912 (Rietz 1912) and entitled Ecological Relations of the Crawfishes of Illinois added to the list of Illinois species O. indianensis, C. robustus (also listed for Illinois by Faxon 1885), C. tenebrosus (as C. bar- toni brevis), and Fallicambarus fodiens This papti is published by aiilhoriiy of the .Stale of Illinois and is a contribution of the Ser- lion of Kaunistif .Surveys and Insect Identification of the Illinois Natural History Survey. Dr. Lav\- rence M. Pa^e is a Zoologist in the- .Sec tion. (as C. argillicola, also listed for Illinois by Faxon 1890). A doctoral dissertation submitted to the University of Illinois in 1955, The Biology of the Crayfishes of Central and Southeastern Illinois (Brown 1955), was based on extensive collecting (at 4 10 stations) in the Sangamon, Wabash, and Table 1.—Classification of the crayfishes and shrimps of Illinois, primarily following Holthuis (1952) and Hobbs (1974b). Order Decapoda Family Palaeinonidae Genus Macrobrachiurn 1. M. ohione (Smith) Genus Palaemonetes 2. P. kadiakensis Rathbun Family Cambaridae Subfamily Cambarellinae Genus Canibarellus Subgenus Dirigicambarus 3. C. shufeldlii (Faxon) Subgenus Pandicambarus 4. C. puer Hobbs Subfamily Cambarinae Genus Procambarus Subgenus Girardietla 5. P. gracilis (Bundy) Subgenus Ortmannicus 6. P. acutus (Girard) 7. P. viaeinridis (Faxon) Subgenus Scapulicambarus 8. P. clarkii (Girard) Genus Orconectes 9. O. illinoiensis Brown 10. O. tmniunis (Hagen) 11. O. indianensis (Hay) 12. O. kentuckiensis Rhoades 13. O. lancifer (Hagen) 14. O. placidus (Hagen) 15. O. propinquus (Girard) 16. O. rusticus (Girard) 17. O. stannardi Page 18. O virilis (Hagen) Genus Fallicambarus Subgenus Creaserinus 19. F. fodiens (Cottle) Genus Cambarus Subgenus Erebicambarus 20. (.'. tenebrosus Hay 21. C rusticijormis Rhoades Subgenus Lacunicambarus 22. (.'. diogenes Giraid •Subgenus Puncticamhnrus 23. (',. robuslus Girard 335 336 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 Ohio river systems. To the growing list of lUinois species, including Cambar- ellus shujeldtii and O. lancijer reported in the interim by Faxon (1914), Brown added P. clarkii, O. kentuckiensis, and O. illinoiensis (described by Brown in 1956). Statewide collections made for the present survey have added Cambarellus puer, P. viaeviridis, O. placidus, O. sta?}nardi, and Cambarus rusticiformis to the list of Illinois species. The list of Illinois shrimps and crayfishes now stands at 23 species. METHODS Collecting for the present survey of Illinois decapods spanned a decade (March 1972 to September 1982). Col- lections were made at 1,294 localities (Fig. 1) in all counties of the state (Fig. 2), but were most heavily concentrated in southern Illinois, where the largest diversities of habitats and species occur. All species of aquatic Malacostraca (Isopoda, Amphipoda, Decapoda) present at each site were collected. A plethora of taxonomic problems among the isopods and amphipods prevent their inclusion in this report. Lewis & Bowman (1981) and Lewis (1982) re- cently have published distributional studies on the subterranean isopods of Illinois. A dot on Figure 1 indicates that one or more malacostracans were collected at that locality. Stations sampled at which no species were found are not shown on the map. Although the ab- sence of malacostracans at a site can be informative, it may indicate only that insufficient time was spent in searching. Old records (pre- 1972) are included on the species distribution maps when they document significant changes in distri- bution (e.g., for Mac robrachiuin ohione). A lew recently (1984) dis- covered localities are plotted for O. rusticus and O. stannardi. Because ecological analyses were attempted for each species, all indi- viduals encountered usually were pre- served; if large numbers were encoun- tered, only the early portion of the sample was preserved, and subsequently collected individuals were returned to the water. Specimens usually were collected by dipnetting, minnow seining, or by digging them from their burrows. Malacostraca require cover, and the most successful method of collecting was to place a dip net downstream from, or (in standing water) next to, accumu- lations of stones, brush, living vegeta- tion, etc., and to kick through the material, thereby dislodging specimens into the dip net. The most successful method of collecting burrowers was to dig into the burrow down to the water table, splash the water, wait for the curious crayfish to investigate the dis- turbance, and grab it. This worked well in the relatively shallow burrows of C. diogenes but less well in the deepjer burrows of P. gracilis. P. viaei'iridis, and F. fodiens. Specimens were placed in 10-percent formalin if large, in 70-percent ethyl alcohol if small enough to go into a 3-dram \ial. In the laboratory, speci- mens were washed in water, identified, counted, sexed, and placed in 70- percent ethyl alcohol for permanent storage in the Illinois Natural History Survey collection. Records were kept on the presence of form I males and of females carrying eggs or young. Measurements of crayfishes are given in millimeters of carapace length (CL). Length-frequency distributions are presented for those species for which no published growth or longevity data are available and of which large col- lections were made in Illinois. Keys, diagnoses, and descriptions are based on Illinois populations only. Taxa are arranged phylogenetically. except within Orcoiifi tes where species arc arranged alphabeticalh because of our present lack of understanding of inter- specific relationships. EXTRALIMITAL SPECIES AND STUDIES Several spec ies of crav fishes not found August 1985 Crayfishes and Shrimps of Illinois 337 Fig. 1.— Localities in Illinois where aquatic Malacostraca were collected between March 1972 and September 1982. 338 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 in Illinois live in eastern Missouri and western Kentucky in streams draining directly into the Mississippi and Ohio rivers, and they eventually may be found, presumably as waifs, in Illinois. In Missouri, these are Orconectes har- risoni, O. hylas, O. luteus, O. medius, O. nais, O. peruncus, P. punctimanus, O. quadruncus, and Cambarus hubbsi, and in Kentucky, O. bisectus and O. tricuspis. It is more likely, however, that additions to the Illinois fauna will result from human introductions, deliberate or otherwise, and could originate from anywhere. Regional publications on freshwater decapods of North America are avail- able for Alabama (Bouchard 1976), STEPHENSON WINNEBAGO BOONE McHENRt Fig. 2—Coun- ties of Illinois. August 1985 Crayfishes and Shrimps of Illinois 339 Arkansas (Bouchard & Robison 1980), California (Riegel 1959), Colorado (Unger 1978), Florida (Hobbs 1942), Georgia (Hobbs 1981), Indiana (Hay 1896; Eberly 1955), Iowa (Phillips 1980), Kansas (Williams & Leonard 1952), Kentucky (Rhoades 1944a), Louisiana (Penn 1952, 1956, 1959; Penn & Marlow 1959), Maryland (Meredith &: Schwartz I960), Michigan (Pearse 1910; Creaser 1931), Nebraska (Engle 1926), New Jersey (Fowler 1912; Francois 1959), New York (Crocker 1957), Ohio (Turner 1926; Rhoades 1944b), Oklahoma (Creaser & Ortenburger 1933; Reimer 1969), Ontario (Crocker & Barr 1968), Pennsylvania (Ortmann 1906), Texas (Penn &: Hobbs 1958), West Virginia- (Newcombe 1929), Wisconsin (Creaser 1932), the Cumberland Plateau and Cumberland Mountains (Bouchard 1976), New England (Crocker 1979), the Ozark Plateaus and Ouachita Provinces (Williams 1954), the Southern Appa- lachians and Cumberland Plateau (Ortmann 1931), and Mexico (Villa- lobos 1983). Extremely useful publications on crayfishes are Hobbs' key and check- list to all described crayfishes of North America (Hobbs 1972a, 1974b). GEOLOGICAL HISTORY OF ILLINOIS AND ZOOGEOGRAPHY OF NATIVE ILLINOIS DECAPODS Prior to settlement by Europeans, Illinois was covered by oak-hickory forest, maple-basswood forest, and bluestem prairie. Today much of Illi- nois has been cleared and developed, primarily for agriculture and urbaniza- tion. The oldest prominent physiographic features of Illinois are remnants of the Appalachian Revolution near the close of the Paleozoic. The intense folding and faulting during the formation of the Appalachian Mountains, and the more moderate folding and tilting to the west, created the template for the topographic diversity present today in southern Illinois (Fenneman 1938). During the Mesozoic, the seas enlarged, and by the Cretaceous they inundated the Coastal Plain as far north as south- ern Illinois. The subsequent uplifting of the continent in the late Cretaceous exposed much of the Coastal Plain, in- cluding the flatlands persisting today as swamps along the lower Ohio River in southern Illinois (Fenneman 1938). In the late Tertiary the Mississippi River ran along the western edge, and the lowermost Ohio River followed the eastern edge, of Crowley's Ridge (a ridge on the Coastal Plain of south- eastern Missouri and northeastern Arkansas). The two great rivers met near where Helena, Arkansas, is now (Fenne- man 1938; Fisk 1944). The Ohio River was comparatively small, had its head- waters in southern Indiana and central Kentucky (Wayne 1952), meandered across southern Illinois through the present Cache River valley, and, after being joined by the Cumberland and Tennessee rivers, flowed south to its confluence with the Mississippi. It is uncertain whether the Tennessee River joined the Ohio above the entrance of the Ohip into the Cache lowland or whether it flowed from its present mouth down the present course of the Ohio and Mississippi rivers and joined the Ohio near southern Tennessee. The Mississippi River followed much of its present course in Minnesota and Wisconsin, then cut through northern Illinois (Willman & Frye 1970), where it was joined by its major tributary, Teays River, and flowed south along the present course of the Illinois River. The Teays River began along the western side of the Blue Ridge Mountains of Kentucky, Virginia, and West Virginia, flowed northward to central Ohio, then turned west across central Indiana and Illinois through the Mahomet Valley to the Mississippi (Horberg 1945; Wayne 1952; Teller 1973); or in Indiana turned south through the present Wabash Valley to the Ohio River (Fidlar 1948). It may be that an early Pleistocene ice advance blocked the original course of 340 Illinois Natural History Survey Bulletin Vol. 33, An. 4 the Teays through the Mahomet Valley and diverted it into the Wabash Valley (Wayne 1952). In the Pleistocene, continental ice caps advanced in stages and interrupted the progress of post-Tertiary erosion cycles. In glaciated areas, topographic diversity was reduced, valleys were ob- structed, and river courses were altered. Areas of low relief, including the Cen- tral Lowlands, were covered by deep deposits of drift, forming undulating till plains. The earliest glacial invasions, the Nebraskan and the Kansan, extended into northern Missouri, southern Il- linois, southern Indiana, and south- central Ohio. Streams which formerly had drained northward into Hudson Bay and eastward to the Atlantic Ocean were blocked and diverted to the Mis- sissippi basin. The lower Teays River was eliminated, and its tributaries were diverted into the ancestral Ohio River (Flint 1971; Hocutt, Denoncourt, Sc Stauffer 1978), greatly increasing the area drained by the Ohio. To the west of the Mississippi River the Kansan ice cap extended farther south than other Pleistocene glaciers and terminated in Missouri, approxi- mately along the present course of the Missouri River. East of the Mississippi River the Kansan ice cap was exceeded in its southern extent by the later Illi- noian glacier, which in some areas reached the present course of the Ohio River. The Illinoian leveled or buried in drift most of Illinois. Because it was the last of the glaciers, drift of the Wis- consinan covers a large area of Illinois (Fig. 3) and its ecological impact is enormous. As the ice caps retreated, huge quan- tities of drift were left behind, filling river channels and valleys. Mounds of drift (moraines) sometimes reached heights of 30 m or more and widths of several kilometers. Moraines generally were concentric to one another and, when they formed extensive lakes, lacked drainage outlets. Ultimately, low COASTAL PLAIN Fig. 3.—Glacial boundaries and major ptiys- iographic features in Illinois. points of the moraines became drainage outlets. Water flowing from one lake to another along the concentric mo- raines gradually formed streams and, eventually, complete drainage systems. With drainage, the process of erosion began. As sediments were carried into lakes, the coarser particles (sand and gravel) were deposited and the smaller particles (silt and clay) were carried away. Eventually, the lakes became marshes and. later, prairies. Thick layers of sand were sometimes left be- hind, such as those found today in the Kankakee Ri\er system of Illinois and Indiana. Dunes were subsequently formed as the wind piled sand into hills. Throughout the Pleistocene, an area of southwestern Wisconsin and north- western Illinois remained imglaciated. August 1985 Crayfishes and Shrimps of Illinois 341 This "Driftless Area" supports a topo- graphic diversity, due in part to recent erosion, absent in the glaciated region. Interglacial periods, the Aftonian, Yarmouth, and Sangamon, were char- acterized by warmer and drier climates similar to that of today (Braun 1950). The Wisconsinan ice cap was up to 3 km thick and in some areas, melting, interrupted by periodic advancement, took more than 10,000 years (Clark & Stearn 1960). New drainage systems eroded the glaciated area, and the junc- tion of the Ohio and Mississippi rivers moved progressively northward (Mat- thes 1933; Fisk 1944). The Great Lakes were formed as water was impounded ip the former Laurentian River basin. The present drainages of Illinois (Fig. 4) are discussed in detail by Forbes & Richard- son (1908) and Smith (1971). Pleistocene glaciers that covered most of Illinois had profound effects on the distributions of organisms. The princi- pal zoogeographic effects were the elimination of populations (probably including entire species) and the dis- placement of species farther south than they had occurred preglacially. Princi- pal postglacial zoogeographic events were the reinvasions of glaciated regions and the crossing of previously insur- mountable basin divides through the newly established drainage systems. The Illinoian glaciation reached within 30 km of the northern boundary of the Mississippi Embayment in south- ern Illinois (Willman & Frye 1970). A few species may have maintained pop- ulations in the surviving uplands, the Shawnee Hills of southern Illinois (Fig. 3). It must have been easy for animals to invade the new drainages of the gla- ciated region of Illinois as they became connected to the existing Ohio and Mississippi rivers. Postglacial invasions (cjr reinvasions) undoubtedly were mostly from the south; i.e., the Ozark Plateau, Mississippi Embayment, In- terior Lowland, Appalachian Plateau, and ephemeral drainages along the southern edges of the glaciers. Because all of northern North America (48° and north) except a large part of Alaska was covered by ice, the only possible non- southern postglacial origins of the decapod fauna of Illinois are east coast drainages (presumably via the Sus- quehanna Outlet connecting the eastern Great Lakes and the Susquehanna River), the Driftless Area of north- western Illinois, and the upper Missouri River drainage. Some species may have invaded from more than one refugium. Species likely, because of their present distributions, to have invaded from the south are M. ohione, P. kadiakensis, P. acutus, F. fodiens, and C. diogenes. All but C. diogenes are sluggish-water inhabitants that probably invaded the low-gradient drainages of Illinois from similar low-gradient streams on the former Mississippi Embayment. C. diogenes presumably moved from the Shawnee Hills and other southern areas into new drainages to the north. P. gracilis, O. im?nunis, andO. I'irilis now occupy western drainages that were unglaciated, principally the upper Missouri, and they probably entered Illinois from the west. The eastward movement of other grassland and plains animals during the recent Xerotliermic Period is discussed by Smith (1957). O. propinquus now occurs only in glaciated areas of eastern North America but is most closely related to species living in streams in unglaciated south- ern Ohio, eastern Kentucky, West V'ir- ginia, and Pennsylvania (Fitzpatrick 1967). It and C. robustus probably in- vaded Illinois from the east. C. robustus, as discussed in the species account, has since retreated from Illinois. O. stannardi does not now occur in areas considered to have been Pleis- tocene refugia, and it may i)e auto- chthoncjus to the Little Wabash River system of Illinois. Autochlhonism would require it to have differentiated from its closest relatives since the Pleistocene. The present decapod fauna of Illinois 342 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 Michigan Fig. 4.— Present drainages of Illinois. August 1985 Crayfishes and Shrimps of Illinois 343 Table 2.—Native species grouped according to their distributions among physiographic divisions of Illinois. Coastal Plain Shawnee Hills Lowland Glaciated Area Statewide C. puer C. shujeldtii P. clarkii P. viaeinridis O. tancifer O. itlinoiensis O. indianensis O. kentuckiensis O. placidus C. tenebrosus" M. ohione P. kadiakensis P. gracilis O. propmquui O. slannardi O. virilis' C. robuslus P. acutus O. immunis F. jodiens C. diogenes "Also found on the Coastal Plain. .\lso found in one locality in east-central Illinois. may be divided into Coastal Plain, Shawnee Hills, Lowland, Glaciated- area, and Statewide species groups (Table 2). The Statewide species group includes two burrowers (C. diogenes and F. jodiens) and two sluggish-pool inhabitants {P. acutus and O. im- munis). Those sfjecies found on the Coastal Plain and in the Shawnee Hills pre- sumably have been there for a long time and have undergone little or no post- glacial dispersal. Those in the glaciated regions must have invaded or re-invaded after glaciation (i.e., within the past 10,000-100,000 years) or must have speciated since then. Illinois' endemic decapods, O. illtnoiensis and O. slan- nardi, and its near-endemics, O. ken- tuckiensis and O. indianensis, occur south of the boundary of the Wiscon- sinan glaciation. All except O. slan- nardi, which occurs on Illinoian glacial till, are occupants of the Shawnee Hills. NATURAL HISTORY Crayfish Life Cycle During the mating season, form I males (males in which one or more of the terminal elements on the gonopods [first pleopods] are corneous) actively seek receptive females by grasping other crayfishes with their chelae. When a male or nonreceptive female is grasped, it resists, and the aggressive male re- leases his catch and continues search- ing. When a receptive female is grasped, it stretches the antennae, chelae, and walking legs forward, curves the end of the abdomen ventrally and anteriorly. and becomes motionless. The male turns the female on her back, holds her chelae in his, positions his abdomen over hers (still curved forward), inserts the ischial hooks on his pereiopods into the coxal membranes of the pereiopods of the female (which locks the two cray- fish together), and inserts the tips of his gonopods in the annulus ventralis (seminal receptacle) of the female. Sperm travel from the reproductive openings on the bases of the fifth pe- reiopods along the gonopods to the annulus ventralis, and mating is com- pleted. A waxy sperm plug projecting from the annulus indicates that the female has recently mated. Frequencies of form I males among Illinois collections of crayfishes indicate that most matings occur between Sep- tember and March, although consider- able interspecific variation occurs. Species of Cambarellus have a high incidence of form I males throughout the year. Sperm are carried by the female until oviposition (egg laying), which in Illinois occurs mostly in March, April, and May but may begin in some species as early as December or January. In preparation for oviposition the female cleans the underside of her abdomen, mostly through brushing motions of the fifth pereiopods. During oviposi- tion the female turns on her back, curls the abdomen forward, and from glands on the abdomen secretes a clear, sticky substance (glair) which fills the area between the curled abdomen and the thorax. Sperm and eggs are released 344 Illinois Natural History Survey Bulletin Vol. 33, An. 4 Fig. 5.—Female Orconectes immunis with eggs attached ("in berry"). into the glair from the anmilus ventrahs and external openings of the oviducts, respectively. Following oviposition, the glair hardens and cements the fer- tilized eggs to the pleopods. With eggs attached to her abdomen, a female cray- fish is said to be ovigerous or "in berry" (Fig. 5). Eggs are carried by the female for 2-20 weeks, depending on the water temperature. When the eggs hatch, the young remain attached to the abdomen and are carried by the female througfi three instars. The first juvenile instar has extremely large eyes, a yolk-filled carapace, an incomplete abdomen, and hooked chelae. First instars attach to the female with the hooked chelae and by a stalk linking the telson of the instar to the abdomen of the mother. After 2-7 days first instars molt to second instars, which are more adultlike but still lack a fully developed abdomen and cling to the mother's abdomen by their chelae. After another 4-12 days, another molt results in the third instar, which closely resembles the adidt and hangs onto the mother with its chelae and pcreiopods. It may leave the protec- tion of the mother for short periods. Subsequent instars are free living. Growth proceeds through a series of (i-10 molls during the summer, and sexual maturity may be attained by late summer or early fall. Sexual matu- rity in the male corresponds to a molt to form I. The first molt following the mating period returns the male to a nonieproducti\e state (form II) in which the gonopods are without cor- neous tips. Following copulation, females do not molt until after the young permanently have left. In Illi- nois, crayfishes appear to survive a maximum of two reproductive seasons. In nonburrowing Illinois crayfishes, all aspects of their life history occur in surface waters. Burrowing species spend variable amounts of time in surface waters. Adults of P. gracilis and P. I'iaeviridis leave their burrows and enter temporary bodies of v\ater to mate. F. fodiens leaves its burrows to mate, and females remain out of their burrows during the period in which they carry their eggs and young. Young of these three species remain above ground in temporary water as long as the water remains; as the water table recedes below the stnface, they burrow, .\dults of P. I'laei'iridis and F. fodiens may be foimd above ground in Illinois onl\ during periods of late winter-to-spriiig flood- ing (January to May). Sexually mature P. gracilis leav e bmrows on warm lainv nights but otherwise seldom are found out of their burrows, .\dults of C. dio- genes leave their burrows on the banks August 1985 Crayfishes and Shrimps of Illinois 345 of streams more often than do the species just mentioned, perhaps to forage as well as to mate, but do so more frequently during the mating and egg- and young-carrying seasons. Freshwater Shrimp Life Cycle Females carry eggs attached to their pleopods between April and August in Illinois and between February and October farther south. The eggs hatch, and free-swimming larvae pass through six stages in about 3 weeks, described in detail for Palaemonetes kadiakensis by Broad &: Hubschman (1963). Larvae and juveniles increase, and adults de- crease, in abundance through the sum- mer. Adults disappear in late summer- early fall, corresponding to a maximum life span of about 1 year in Palaemo- netes kadiakensis and 2 years in Macro- brachium ohione. The largest adults mature and reproduce earliest in the year. Observations on copulation ap- parently are unrecorded. KEY TO ILLINOIS SPECIES Morphological characteristics used shrimps in keys and systematic accounts to describe and identify crayfishes and are illustrated in Fig. 6-9. cephalic process central projection caudal process mesial process A central projectioi mesial process B Fig. 6.—Lateral view of Macrobrachium ohione (after Hoitfiuis 1952). Fig. 7.—Morpfiology of crayfish gonopods (after Hobbs 1972a). (A) Mesial (left) and lateral views of terminal elements of generalized Pro- cambarus gonopod. (B) IVIesial (left) and lat- eral views of terminal elements of generalized Cambarus gonopod. (C) Generalized Orco- nectes gonopods. Crayfishes are most easily and accu- rately identified by using characteristics of the gonopod (first pleopod) of form I (breeding) males, and in geographic areas having many similar species, accurate identifications of specimens f)ther than form I males can be nearly impossible. In Illinois, with only 23 species of decapods, it is possible to identify most free-living specimens (including juveniles) to species. Fol- lowing are two keys, the first is based on general morphology, the second on ( harac terisiif s of form I male cray- fishes. In both keys, references to cray- fish gonopods are to those of form I males. 346 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 antennule antenna antennal scale acumen of rostrum marginal spine - rostrum suborbital angle postorbital ridge hepatic spine cervical spine length of carapace — width of areola length of areola — uropod Fig. 8— Dorsal view of crayfish, illustrating features used in keys and descriptions (after Hobbs 1972a). August 1985 Crayfishes and Shrimps of Illinois 347 dactyl length of palm hook on ischium pleopods I width of palm Fig 9—Ventral view of crayfish, Illustrating features used in keys and descriptions (after Hobbs 1972a). 348 Illinois Natural History Survey Bulletin Vol. 33, An. 4 I. Key to Decapods of Illinois (Key to form I male crayfishes on page 35i) 1. First two pair of legs with chelae; abdomen compressed laterally (Fig. 6)... (Family Palae- monidae) 2 1. First three pairs of legs with chelae; abdomen flattened dorsoventrally (Fig. 8)... (Family Cambaridae) 3 2. Second pair of legs much longer than the first pair; 9-13 teeth along upper edge of rostrum (Fig. 10) Macrobrachium oluone (p. 356) 2. Second pair of legs only slightly longer than first pair; 6-8 teeth along upper edge of rostrum (Fig. 11) Palaernoyieles kadiakensis (p. 359) 3. Areola wide; narrowest width about 20-25 per- cent of length (Fig. 12); rostrum flat dorsally; small, adult carapace length never more than 16 mm (32 mm total length). . .(Genus Cam- carellus) 4 Areola obliterated to moderately wide but nar- rowest width never more than 20 percent of length (except in Orconectes propinquus and O. stannardi); rostrum conca\e dorsally; maxi- mum carapace length much more than 16 mm 5 4. Terminal elements of gonopod cur\ed (Fig. 13); margins of rostrum slightly convex, barely con- verging anteriorly; annulus vcntralis extends anteioventialh (Fig. 14) Catnbarellus jjury (p. 366) 4. Terminal elements of gonopod straight (Fig. 15); margins of rostrum straight, strongly con- verging anteriorly; annulus ventralis extends ventrallv (Fig. 16) Cambarrllus shufrldtii (p. ;i63) Fig. 10. Fig. 11. Fig. 12. c Fig. 13. Fig. 14. Fig. 15. ./^ Fig. 16. August 1985 Crayfishes and Shrimps of Illinois 349 5. AiTola linear to obliterated (Fig. 17) 6 5. Areola narrow to wide 10 6. Rostrum deeply excavated, with acumen longer than basal margin of rostrum (Fig. 18) Orconectes lancijer (p. 400) 6. Rostrum shalknvly excavated; acumen much shorter than margin of rostrum 7 7. Chela long and narrow, maximum width less than one-third of length (Fig. 19); acumen and marginal spines of rostrum well-developed (Fig. 20); a longitudinal blue stripe along underside of abdomen (fades in preservative) Pro- cambarus clarkii (p. 381) Fig. 17. Fig. 18. Fig. 19. Fig. 20. 7. Chela wide, maximum width more than one- third of length; small acumen, no marginal spines on rostrum; no blue stripe on abdo- men 8 1!^.. w Fig. 22. Fig. 21. 8. Areola linear only at midlength (Fig. 21); gono- pod terminates in small straight elements (Fig. 22); annulus ventralis subrectangular, elevated laterally Procambarus gracilis (p. 370) 8. Areola linear through most of length (Fig. 23); gonopod terminates in two large curved ele- ments (Fig. 24); annulus ventralis wider than long, elevated posteriorly 9 Fig. 24. 350 Illinois Natural History Survey Bulletin \'o1. 33. An. 4 9. Suborbital margin of carapace smooth to slight- ^^^ ly angular (Fig. 25); opposable margin of mov- ^^ y^ able finger of chela with deep concavity (absent ^ \r~^ '^'^ ^^ in regenerated chela) in proximal one-half (Fig. 26) Fallicambarus jodiens (p. 422) Fig. 26. 9. Suborbital margin of carapace angular (Fig. 27); opposable margin of dactyl of chela without ^:^^^^^^^ Z^ deep concavity (Fig. 28) Cambarus y^ PI 27 diogenes (p. 434) ^\^ Fig 28. Fig. 29. 10. Margins of rostrum without large spines (Fig. 29) 11 10. Margins of rostrum with large spines (Fig. 30) 15 11. Carapace laterally compressed: chela long and narrow, maximum width less than one-third of length 12 11. Carapace dorsoventrally flattened; chela wide, maximum width more than one-third of length Il4 12. Gonopod terminates in large curved elements (Fig. 31); annulus \entralis wide, with fossa far to one side (Fig. 32) Orconectes immunis (p. 388) 12. Gonopod lerniinates in very short elements; Fig. 31. annulus veniralis with fossa more centrally located 13 Fig. 32. Fig. 30. i August 1985 Crayfishes and Shrimps of Illinois 351 13. Margins of rostrum ele\ated, strongly converg- ing to acumen (Fig. 33); narrow areola (Fig. 33) Procambarus acutiis (p. 376) 13. Margins of rostrum not elevated; rostrum more rounded anteriorly (Fig. 34); wide areola (Fig. 34) Procambarus viaeviridis (p. 378) 14. Base of fixedfingerof chela with deep dorsal and ventral impressions (Fig. 35); margins of ros- trum converge anteriorly (Fig. 36) Cam- barus robustus (p. 439) 14. Base of fixed finger of chela without deep im- pressions (Fig. 37); margins of rostrum nearly parallel, barely converging anteriorly (Fig. 38) Cambarus tenebrosus (p. 428) 15. Margins of rostrum distinctly concave (Fig. 39) ^ 16 15. Margins of rostrum straight (Fig. 40) 18 16. Central projection of gonopod large and blade- like, curved at right angle to shaft (Fig. 41); annulus ventralis with fossa to side Cam- barus rusltcijortnis (p. 432) 16. Clentral projection of gonopod not large and bladelike; annulus ventralis with fossa centrally located 17 17. Dorsal (cephalic when lifted from body) surface of gonopod with prominent shoulder (Fig. 42) Orconectes ruslicus (p. 412) Fig. 33, Fig. 34, Fig. 38. Fig. 37. Fig. 40. Fig. 39 Fig. 41 Fig. 42. 352 Illinois Natural History Survey Bulletin Vol.33. Art. 4 17. Dorsal surface of gonopod without prominent shoulder (Fig. 43) Orconectes placidus (p. 404) 18. Areola distinctly narrows just anterior to mid- length (Fig. 44); gonopod reaches coxa of cheli- ped with abdomen flexed; no black band on fingers of chela Orconectes virilis (y). AM) ^b^^j^ ! Fig 43. 18. Areola remains fairly wide throughout (Fig. 40); Fig. 44. gonopod does not reach coxa of first pair of walking legs; black band (fades in preservative) near tips of fingers of chela 19 19. Rostrum deeply excavated medially, with mar- gins distinctly converging anteriorly (Fig. 45) Orconectes illinoiensis (p. 386) 19. Rostrum shallowly excavated, margins barely p| ^^ converging anteriorly 20 20. Rostrum without median carina (Fig. 46);. Orconectes kentuckiensis (p. 396) 20. Rostrum with median carina (small in O. in- dianensis) (Fig. 47); 21 21 . Mesial process of gonopod with spur on caudal surface (Fig. 48) Orconectes stnnnardi (p. 415) 21. Mesial process of gonopod without spur 22 22. Rostrum with large median carina (Fig. 49); gonopod with tips of elements barely di\erging Fig. 49. (Fig. 50); Orconectes propinquus (p. 406) Fig 48. / Fig. 47. Fig. 50. / 22. Rosinun with small meilian carina (Fig. 51); p| ^^ gonopod with tips of elements stroiigh di\erg- ing (Fig. 52).... Orcuneclrs indiiinensis (p. 394) Fig. 52. August 1985 Crayfishes and Shrimps of Illinois 353 II. Key to Illinois Crayfishes Based on Form I Males (i.e., males with corneous ter- minal elements on the gonopods) 1. Ischia of second and third peieiopods with hooks (Fig. 9). . .Genus Cambarellus 2 1. Ischia of second pereiopods without hooks ....3 2. Terminal elements of gonopod straight (Fig. 53) Cambarellus shufeldtii (p. 363) 2. Terminal elements of gonopod curved at about right angle to shaft (Fig. 54) Cambarel- ^'^^ 53. Fig 54 lus puer (p. 366) 3. Central projection of gonopod large and blade- like, curved at right angle to shaft (Fig. 55). . . Genera Fallicambarus and Cambarus 4 2. Central projection of gonopod not bladelike. . . Genera Orconecles and Procambarus 8 Fig. 55 4. Areola linear (Fig. 56) 5 ^ * 1 1 ^ Fig. 56. 4. Areola narrow to wide 6 5. Suborbital margin of carapace smooth to slight- ly angular (Fig. 57); opposable margin of mov- able finger of chela with deep concavity (absent Fig. 57. in regenerated chela) in proximal one-half (Fig. 58) Fallicambarus fodiens (p. 422) Fig. 58. Suborbital margin of carapace angular (Fig. 59); '^ ^ opposable margin of movable finger of chela without deep concavity (Fig. 60) Cam- barus diogenes (p. 434) Base of fixed finger of chela wiili dccj) dorsal and ventral imjiressions (Fig. 61) Cam- barus robuslus (p. 439) "^'9 ^1 354 Illinois Natural History Survey Bulletin Vol. 33, An. 4 6. Base of fixed finger of chela without deep im- pressions 7 7. Margins of rostrum concave, with spines (Fig. 62) Cambarus rusticiformis (p. 432) 7. Margins of rostrum straight, without spines (Fig. 63) Cambarus tenebrosus (p. 428) 8. Gonopod terminates in two elements (Fig. 64) . . .Genus Orconectes 9 8. Gonopod terminates in more than two, very short elements (Fig. 65). . .Genus Procambarus 18 Fig. 62 Fig. 63. 9. Areola obliterated (Fig. 66); rostrum deeply H / excavated, with acumen longer than basal ^V ' margin of rostrum (Fig. 66) Orconectes lancifer (p. 400) 9. Areola narrow to wide; acumen much shorter than basal margin of rostrum 10 ^^9- 65. / Fig. 66. 10. Both terminal elements of gonopod curved at right angle to shaft (Fig. 67); annulus ventralis with fossa far to one side (Fig. 68) Orco- nectes inimunis (p. 388) 10. One or both terminal elements of gonopod ^ft '^'9 ^8. straight or only slightly curved; annulus ven- tralis with fossa more centrally located 11 Fig. 67. 11. Central projection constitutes one-fourth or less of total length of gonopod (Fig. 7) 12 11. Central projection constitutes more than one- fourth of total length of gonopod (Fig. 7) ...16 12. Terminal elements of gonopod distinctly diver- gent (Fig. 69 & 70) 13 Fig.'69. 72. August 1985 Crayfishes and Shrimps of Illinois 355 12. Terminal ek-ments of gonopod nearly parallel (Fig. 72 & 73) .'. 14 u // 13. Terminal elements of gonopod curved (Fig. 69) Orconectes kentuckiensis (p. 396) 13. Terminal elements of gonopod nearly straight (Fig. 70) Orconectes indlauensis (p. 394) 14. Rostrum with median carina (Fig. 71) 15 14. Rostrum without median carina Orco- nectes illinoiensis (p. 386) Fig. 71 15. Mesial process of gonopod with spur on caudal surface (Fig. 72) Orconectes stannardi (p. 415) 15. Mesial process of gonopod without spur (Fig. 73) Orconectes propinquus (p. 406) M p| 73 16. Both elements of gonopod curved (Fig. 74); areola distinctly narrows anteriorly (Fig. 75); no black bands on fingers of chela Orco- nectes I'irilis (p. 417) Fig. 74. 16. Mesial process of gonopod not curved; areola remains fairly wide throughout; black, band (fades in preservative) near tips of fingers of chela 17 Fig. 75. Fig. 76. 17. Dorsal (cephalic when lifted from body) surface of gonopod with prominent shoulder (Fig. 76) Orconectes rusticus (p. 412) 17. Dorsal surface of gonopod without prominent shoulder (Fig. 77) Orconectes placidus (p. 404) 18. Dorsal surface of gonopod with prominent shoulder (Fig. 78); cephalic process a large lobe (Fig. 78) Procambarus clarkii (p. 381) 18. Dorsal surface of gonopod without shoulder; cephalic process small 19 Fig. 78. Fig. 77. 356 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 19. Areola linear at midlength (Fig. 79); hooks on ischia of third pereiopods only (Fig. 9) Procambarus gracilis (p. 370) 19. Areola narrow but not linear (Fig. 80 & 81); hooks on ischia of third and fourth pereio- pods 20 20. Margins of rostrum elevated, converging to acumen (Fig. 80); narrow areola (Fig. 80) Procambarus acutus (p. 376) 20. Margins of rostrum not elevated; rostrum more rounded anteriorly (Fig. 81); wide areola (Fig. 81) Procambarus viaeviridis (p. 378) Fig. 79 Fig. 80. Fig. 81. SYSTEMATIC ACCOUNTS FAMILY PALAEMONIDAE Both shrimps native to Illinois are members of the family Palaemonidae, which is worldwide in distribution. Macrobrachium ohione, with 9-13 teeth along the upper edge of the ros- trum, is easily separable from Palae- monetes kadiakensis, which has only 6-8 (almost always 7) teeth. M. ohione, which reaches 100 mm in total length (tip of rostrum to tip of telson), also is much larger than P. kadiakensis, which reaches a maximum length of only about 50 mm. Genus Macrobrachium Bate Macrobrachium Bate 1868 Species of Macrobrachium, referred to as river shrimps, differ from species of Palaemonetes by possessing a hepatic spine, lacking a branchiostegal spine, and having the second pair of legs much longer than the first pair (in Palae- monetes the second pair is only slighth longer than the first pair). Although Macrobrachium contains aboiii 100 species worldwide (35 in the western hemisphere), only four (A/. ohione, M. acanthurus. M. carcinus, and M. oljersii) are found in the fresh- waters of the eastern United States (Hedgpeth 1949; Holthuis 1952). M. ohione is the only species of Macro- brachium found in the Mississippi River system. Macrobrachium ohione (Smith) (Fig. 82) Palaemon Ohionis Smith 1874 Palaemon sallei Kingsley 1882 Description.—Holthuis (1952) dis- tinguished M. ohione from other American species of Macrobrachium by this combination of characteristics: carpus of second legs as long as or longer than merus; telson with a dis- tinct posterior margin (rather than gradually tapering toward a slender tip); second chelae of adult male equal or nearly equal to one another in size (some species are bilaterally asymmet- rical with the chela on one side much larger); no tubercles along cutting edges of fingers of second chela of adult male; cutting edges of fingers of large chela of adult male with one or two fairly large teeth proximally; eggs num- erous and small (about 0.5 mm in diam- eter); second pair of legs of adult male with velvety pubescence on some or all joints; rostrum with styliform apex (and no teeth distally). M. ohione is pale gray with light blue spots and a blue abdomen (Hedgpeth 1947, 1949). .\duli females average larger than adult males and ha\e much August 1985 Crayfishes and Shrimps of Illinois 357 Fig. 82.—Lateral view of Macrobrachium ohione (after Holthuis 1952). longer second legs (Holthuis 1952). Females reach about 100 mm in length, males about 70 mm (Hedgpeth 1949; Truesdale & Mermilliod 1979). Distribution and Habitat.—M. ohione occupies freshwater habitats along the Atlantic coast from Virginia to Florida, along the Gulf Coast from Alabama to Texas, and north in the Mississippi and Ohio river systems to Oklahoma, Missouri, Illinois, Indiana, Fig. 83.—Total distribution of Macrobrach- ium ohione. and Ohio (Fig. 83). Coastal populations inove into estuaries in spring (Gunter 1937; Hedgpeth 1949; Reimei, Strawn, Sc Dixon 1974). In Illinois, M. ohione has been re- corded from Cairo, Alexander County (Forbes 1876); Grand Tower, Jackson County (Forbes 1876); Chester, Ran- dolph County (McCormick 1934); and Shawneetown, Gallatin County (Hedg- peth 1949) (Fig. 84). Although Hedg- peth (1949) and Holthuis (1952) cite Luce (1933) as recording M. ohione from the lower Kaskaskia River. Illi- nois, the reference is actually to its use as fish bait. Forbes (1876) described A/, ohione as "abundant at Cairo, where it is fre- quently eaten" and reported second handedly its presence in the Mississippi River system from St. Louis to New Orleans. It aj^parently was common in the Mississippi River as recently as the 1930's, when McCormick (1934) described the successful use of "shrimp sets" at Chester. Shrimp sets were wil- low or Cottonwood branches set along the river's edge and bent so that their leaves were under water. As shrimj) fed on the leaves they weie dipneited by fishermen. In the lUintjis Natural History Survey collection are six series of M. ohione from Illinois, including one made by S. A. Forbes at Caircj (undated). The five collections with locality data. 358 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 (Mount Carmel, Wabash County, 1892; Mississippi River, Grand Tower, Jack- son County, 1932 and 1944; Mississippi River, Missouri River Station, Madison County, 1944; and Mississippi River, 1 mi S Cairo, Alexander County, 1962) are represented on the distribution map (Fig. 84). No M. ohione were found during the present survey (1972-1982). Although perhaps still present in the state, the species obviously has under- gone a tremendous reduction in popu- lation. Little information on the ecol- ogy of the species in Illinois is available, but its decline presumably is attribut- able to the loss of suitable habitats as a result of the excessive modifications (especially channelization, impound- ment, and drainage of bottomland lakes) of Illinois' largest rivers. Life History.—In the Mississippi River at Chester, McCormick (1934) Fig. 84 —Distribution of Macrobrachium ohione in Illinois. Large circles = pre-1900 collections, small circles = 1932-1949 collec- tions, black dot = collection made in 1962. counted 8,000 eggs on an 84-min female and observed that ovigerous females ranged from 34 to 90 mm long. Among Illinois Natural History Survey col- lections of Illinois M. ohtone (all made between May and October), ovigerous females are present only in a collection made 13-14 May 1932. In Louisiana, Truesdale Sc Mermil- liod (1979) found ovigerous females (50-93 mm in total length) from March through September. Five females, rang- ing from 54 to 80 mm, were carrying 6,273-24,800 eggs. Eggs were ai)out 0.5 mm across, bright orange when first laid, and brown at later stages. Huner (1977) found M. ohione at Port Allen, Louisiana, to range from 17 to 92 mm long, to average 30.6 mm (N = 7,058). and to live a maximum of 2 years. Ovi- gerous females (N = 88) ranged from 27 to 92 mm and averaged 66 mm. In the springs of 1969-1971, Reimer, Strawn, & Dixon (1974) studied move- ments of M. ohione into Galveston Bay, Texas. Increased movement was associated with rainfall, and salinities as high as 15 percent seemed to prevent further movement. The first individuals to appear in the bay were males and females without eggs. Females carrying eggs first appeared in late March. Fe- males averaged larger than males, and only the largest females (52-82 mm) bore eggs. M. ohione feeds on both plant and animal material (McCormick 1934; Gunter 1937; Darnell 1958; Truesdale & Mermilliod 1979) and in turn is fed upon by several predators, including flathead catfish [Pylodiclis olii'aris) and white bass (Morone chrysops) (Bryan, Truesdale. &: Sabins 1975). Genus Palaemonetes Heller Palaemonetes Heller 1869 Of the approximately 17 species of Palaemonetes (referred to as prawns, glass shrimp, and freshwater shrimp) in the western hemisphere, onlv P. kadiakcnsts occurs in Illinois. P. palu- dosus is widespread in the eastern I'nited States and along the Gulf Coast August 1985 Crayfishes and Shrimps of Illinois 359 Fig. 85— Lateral view of Palaemonetes kadiakensis. as far west as eastern Texas, but it does not enter the Mississippi River system. In addition to P. kadiakensis and P. paludosus, the only freshwater species in the United States are highly localized spring- and cave-inhabiting forms in Texas (P. antrorum, P. holthuisi, and P. texanus) and Florida (P. cummingi) (Strenth 1976). Palaemonetes kadiakensis Rathbun (Fig. 85) Palaemonetes kadiakensis Rathbun 1902 Description.— P. kadiakensis is the only species of Palaemonetes with three apical setae on the appendix masculina (Fig. 86) occurring in the freshwaters of the I'nited States (Fleming 1969). Rostrum straight; upper margin convex with 6-8, usually 7, teeth; lower margin with 2-3 teeth. Carapace with antennal and branchiostegal spines; branch- iostegal spine below branchiostegal groove. Abdomen humped at third segment; sixth abdominal slightly less than twice as long as fifth segment, slightly shorter than telson. Anterior pair of dorsal spines on telson usually distinctly behind middle of length, posterior pair near distal margin. Eyes large, pigmented. Lateral antennular flagellum with rami fused for 11-23 articles; free portion of shorter ramus with 3-8 articles. Scaphocerite about three times as long as wide, lamella overreaching distolateral tooth. Living specimens are transparent with green eyes, red-brown antennae, and many very small red-brown specks on the body. The internal organs are readily visible, and a bright green vege- tation-filled intestine is often a prom- inent feature. The species reaches a Fig. 86.—Appendices masculina and interna (left) and entire second pleopod (right) of male Palaemonetes kadiakensis (after Meehean 1936). 360 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 maximum length of about 53 mm total length (Meehen 1936). Distribution and Habitat. — P. kadia- kensis occupies sluggish heshwater habitats from Minnesota and the shores of Lakes Ontario, Erie, and Michigan south in the Mississippi River system to the Gulf of Mexico, and Gulf Coast drainages from northeastern Mexico (Nuevo Leon) to northern Florida (Hol- thuis 1952) (Fig. 87). Although the Fig. 87.—Total distribution of Palaemo- netes kadiakensis. locality given on the label accompany- ing the type specimen is Kodiak Island. Alaska, it is doubtful that the species ever occurred in Alaska (Holthuis 1952). P. kadiakensis now occurs through- out the southern one-fourth of Illinois, in backwaters along the Mississippi River, in backwaters of the Illinois River at least as far north as Bureau County, and in Wolf Lake (Lake Mich- igan drainage) in Cook County (Fig. 88). It abounds in swamps (Fig. 89) and swamplike streams on the Coastal Plain of Illinois and is common in standing- and sluggish-watei habitats as far north as Salem. It is uncommon in the Illinois River and in Wolf Lake. Fig. 88.— Distribution of Palaemonetes kadiakensis in Illinois. Open circle = pre-1898 record for the Kankakee River; black dots = 1972-1982 collections. Forbes (1876) recorded P. kadiakensis (as P. exilipes) only from the Illinois River but described it as being very (ommon. A specimen in the I'.S. Na- tional Museum of Natural History documents its former occurrence at Kankakee. Kankakee County (pre-1898. exact date unknown), a localitv well outside the present range. P. kadiakensis almost always is asso- ciated with, and is most abundant in. li\ing a(]ualic \egetation. Its reduction in distribution and abundance in Il- linois probably is attributable to in- creased turbidiiv and .sedimentation and the resultant loss of vegetation. Beds of emergent and submerged \ege- t.iiion once were common throughout Illinois but now are rarely encountered. The loss of vegetation is also one of tire August 1985 Crayfishes and Shrimps of Illinois 361 *g*:-HL. Fig. 89.—Mermet Swamp, Massac County, Illinois (8 October 1984). Large populations of Palaemonetes kadiakensis and P. clarkii occur here. main causes of reductions in popula- tions of Illinois fishes (Smith 1979). Life History.— Reproduction in Illinois, as indicated by the presence of ovigerous females, occurs from April to August (Table 3). In Louisiana the reproductive period extends from Feb- ruary to October (Meehean 1936; White 1949), and in central Missouri (where the species is introduced) from mid-May to August (Nielsen k Reynolds 1977). In Illinois collections, ovigerous females range in total length from 30 to 39 mm. Eggs (embryos) on 15 females numbered 22-137 (mean, 86.6) (Table 4). The relationship (Model II regres- sion) between the number of eggs (N) and total length (L) was N = -230.16 + 9.19L, r = 0.674, and between the number of eggs and the weight of the female (W) was N = 10.78 + 318.21W, r = 0.694. Nielsen &: Reynolds (1977) found that the number of eggs varied from 20 to 76 on females 25-36 mm in length and equalled -111.43 + 5.29 L, r- = 0.64. Meehean (1936) found as many as 154 eggs (on a 49-mm female). Eggs are spherical, orange, and about 1.0 mm in diameter. The egg mass con- stituted an average of 11.2 percent of the weight of the female (without the eggs) in 15 Illinois females (Table 4). Females may produce more thair one brood per season (Broad &: Hubschman 1963; Nielsen & Reynolds 1977). The incubation period lasts 24-28 days at 18.5°-24°C. Detailed accounts of lar- Table 3.— Illinois collections of Palae- monetes kadiakensis. 362 Illinois Natural History Survey Billetin Vol. 33. Art. 4 Table 4 —Relationship between size of female and the number and weight of eggs (em- bryos) in Illinois-collected Palaemonetes kadiakensis. August 1985 Crayfishes and Shrimps of Illinois 363 suture; except in some species of FalU- cambarus, the telson always is divided in North American cambarids. Genus Cambarellus Ortmann Cambarellus Ortmann 1905 Gonopod with three terminal ele- ments; central projection not large and bladelike. Ischia of second and third pereiopods of male with hooks. Species of Cambarellus are distrib- uted throughout the southern United States and Mexico, generally in swamps and other standing- or sluggish-water habitats. They are "dwarf" crayfishes, reaching a maximum total length of about 30 mm. Two of the 17 recognized species of Cambarellus occur in south- ern Illinois. Fitzpatrick (1983) revised Cambarellus, proposing the recogni- tion of three subgenera. Cambarellus shufeldtii (Faxon) (Fig. 90) Cambarus Shufeldtii Faxon 1884 Description.—Rostrum broad, flat, often deflecting downward anteriorly; margins straight, strongly converging anteriorly, ending in large spines; acumen moderately large. Carapace compressed, with cervical spines; sub- orbital margin angular. Areola wide, narrowest part about 19-24 percent of length. Chela narrow and long, smooth; palm without tubercles. Form I gono- pod terminates in three elements, all distally directed (not curved): sclerotized central projection, nonsclerotized mesial process, and nonsclerotized caudal process. Dorsal color variable, from rust red to light brown with two dark brown stripes or rows of spots on either side of the areola extending down the abdomen. C. shufeldtii appears to have no ob- viously close relatives, being the only species in the genus with straight ter- minal elements on the gonopod and the only member of the subgenus Dirigi- cambarus (Fitzpatrick 1983). Distribution and Habitat.— C. shufeldtii occupies bodies of sluggish water, mostly on the Coastal Plain, from southern Illinois to southwestern Ala- bama and eastern Texas (Fig. 91). In Illinois, C. shufeldtii is known only from Alexander, Jackson, Massac, Pulaski, and Union counties in both the Mississippi and Ohio basins. Most localities are on the former Mississippi Embayment, but the species also as- cends the lower M ississippi River Valley and the lower Big Muddy River to about Sand Ridge in Jackson County (Fig. 92). C. shufeldtii was first collected in Il- linois at Cairo by Robert Kennicott (Faxon 1914). Brown (1955) found the species in five sloughs and ditches in Alexander and Massac counties. The 28 collections made in the present survey were from cypress swamps, sloughs, and backwaters. C. shufeldtii is especially common in the heavily vegetated LaRue Swamp-Wolf Lake complex in Union County. C. shufeldtii and C. puer have not been found together in Illinois and apparently compete for suitable habi- tats. In Gulf Coast streams C. shufeldtii is supplanting C. puer (see C. puer account). C. shufeldtii was probably more common and widespread in extreme southern Illinois prior to the extensive clearing of swamps and river flood- plains. Brown (1955) mentioned an earlier collection of a "dwarf crayfish" near Shawneetown (Gallatin County) but was unable to document the occur- rence of Cambarellus that far northeast. In view of the occurrence, or former occurrence, of several swamp-inhabit- ing fishes (Fundulus dispar, Elassoma zonatum, Lepomis punclatus, and L. symmetricus [Smith 1979]) that far northeast, or even into the lowlands of the lower Wabash River drainage, the presence of C. shufeldtii or C. puer in the same area is possible. In Louisiana, C. shufeldtii lives in clear, shallow (less than 40 cm deep), permanent, sun-exposed, mud-bot- tomed, vegetated bodies of water (Penn 1950). C. shufeldtii characteristically does not burrow but may survive peri- ods of drought in subterranean (ells (Penn 1950). Apparently the trayfish 364 Illinois Natural History Survey Bulletin Vol. 33, An. 4 i Fig. 90—Cambare//us shufeldtii. A, dorsal view of carapace; B, dorsal view of right chela: C, annulus ventralis; D, mesial viewof gonopod of form I male; E. lateral view of gonopod of form I male; F. lateral view of gonopod of form II male. August 1985 Crayfishes and Shrimps of Illinois 365 Fig. 91.—Total distribution of Cambarellus shufeldtii. excavates a chamber in the mud bottom of the slough or swamp as drying occurs and then seals the top of the chamber to prevent desiccation. Life History.—Form I males are present in Illinois collections (Table 5) during most months and peak in occur- rence in December, February, and March. In Louisiana, Penn (1942, 1950) found form I males in all months except September, with peaks in January, February, and July. Presumably, mating in C. shufeldtii may occur at anytime but peaks in winter in Illinois and in winter and summer in Louisiana. Females carrying eggs were found in Illinois from February to May, and peaked in occurrence in March (Table 5); females carrying young have been found in Illinois in April, June (Table 5), and July (Brown 1955). In Louisiana, females with eggs or young are jjiesenl in every month, with strong peaks in January-Mardi and June-July (Penn 1942, 1950; Lowe 1961), again suggest- ing two peaks of repiodut ti\i' activity in the southern part of the range. Nine Illinois females, ll.tJ-14.2 mm Fig. 92.— Distribution of Cambarellus shu- feldtii in Illinois; black dots = 1972-1982 col- lections. CL, collected in February and March were carrying 47-99 (mean, 80.1) eggs averaging 1.1 mm in diameter. The re- lationship between numbers of eggs (N) and carapace length (CL) is N = —539.0 + 558.9 log CL, r = 0.86. Four females, 12.4-13.5 mm CL, collected in April 1974 were carrying 15-74 (mean, 44.8) young. The decline from an average of 80.1 eggs to an average of 44.8 young suggests a 44 percent mortality during egg and early instar stages. Penn (1942) found the number of eggs or young on a female to increase with the carapace length of the female, with females 8.5 mm averaging 27 offsjiring and those 1 1 mm averaging 54 offsjjring. \n oxer- all average of 34 offspring per female {N = 100) in Louisiana (Penn 1942) is about one-half the average number of ()ns|)riiig (69.2) [jer female (;V = 13) in Ilhncjis; hc)wever, the comparison is 366 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 Table 5.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Cambarellus shufeldtii. August 1985 Crayfishes and Shrimps of Illinois 367 Fig. 9A~-Cambarellus puer A, dorsal view of carapace; B. dorsal view of right chela; C, annulus ventralls; D, mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male 368 Illinois Natural History Survey Bulletin Vol. 33, An. 4 margins convex, barely converging anteriorly, ending in large spines; acumen moderately large. Carapace compressed, with cervical spines; sub- orbital margin angular. Areola wide, narrowest part about 25 percent of length. Chela narrow and long, smooth; palm without tubercles. Form I gono- pod terminates in three elements: sclero- tized central projection, nonsclerotized mesial process, and nonsclerotized caudal process, all curved at sharp angle (about 45°-90°) to principal axis of gonopod. Color as in C. shujeldtii. The closest relatives of C. puer are C. schmitti and C. lesliei (see Fitzpatrick &: Laning 1976), two occupants of the Gulf Coastal Plain from Mississippi to Florida. C. puer, C. schmitti. C. lesliei, C. diminutus, C. blacki, C. ninae, and C. texanus all are members of the subgenus Pandicambarus (Fitz- patrick 1983). Variation in morphological charac- teristics of C. puer was studied by Chambers, Payne, & Kennedy (1979). Three populations were separable from one another but not given taxonomic status. Illinois populations are part of a larger population extending along the Mississippi River and across south- ern Louisiana. In general, individuals in the east are larger than those in the west; females are larger and have shorter, broader chelae and shorter rostra than do males. Distribution and Habitat. — C. puer ranges from southern Illinois and southeastern Missouri to the Gulf Coast of Louisiana and the lower Colorado River drainage of Texas (Fig. 95). All populations are on the Coastal Plain. In Louisiana, C. puer occurs most frequently in shallow (less than 40 cm deep), clear, permanent, sun-exposed, mud-bottomed, vegetated water bodies (Penn 1950). C. puer was first found in Illinois in 1973 (Page & Burr 1973) and now is known fr(jm seven sites in the Missis- sippi and Ohio basins in I'nion, John- son, and Alexander (ouniies (Fig. 96). In Illinois, it occupies cypress swamps (Fig. 97), effluent streams, and lowland areas which probably were swamps prior to being cleared. \\\ but one col- lection from a roadside ditch were made in permanent water bodies, and most specimens were found among living or dead vegetation. C. puer may be more widespread on the Coastal Plain of Il- linois than present records indicate and almost certainly was more widespread and common prior to the drainage of swamps and lowlands. C. puer and C. shujeldtii were col- lected at 23 sites in Illinois, mostly in Coastal Plain swamps, but never were collected together. Competitive exclusion among species of Cambarel- lus has been documented in Gulf Coast areas (Penn & Fitzpatrick 1962. 1963) and apparently is operating in Illinois. C. puer and C. shujeldtii seem unable to coexist and, at least along the Gulf Coast, C. shujeldtii has supplanted C. puer at several localities within historic times (Penn &: Fitzpatrick 1962. 1963). C. shujeldtii was found experi- mentally to be dominant o\er C puer (i.e., C. puer retreats following contact with C. shujeldtii) (Penn &: Fitzpatrick 1963), suggesting that in nature C. shujeldtii agressi\ely displaces C. puer from certain habitats. Life History.—Louisiana data sug- gest IWC3 ieproducti\e periods (winter and summer) (Penn 1950; Black 1966). although form I males can be found in all months (Black 1966). Illinois data are seasonally incomplete (Table 6), but two reproductive peric:)ds. some- what later (i.e.. late winter-spring and fall) than in Louisiana, may also occur in Illinois. In Louisiana, form I males peak in occurrence from December to March (abc^ut 60-95 percent of all males) and from July to .\ugust (about 45-75 percent) (Penn 1950: Black 1966). In the available Illinois collections (Fable 6), form I iiiak's are present February- May and in October. In Louisiana, form I C. puer males measure 7.2-12.0 mm CL and weigh 75.4-412.1 mg (Black 1966). Fhirteen or 14 molts are required to reach sexual maturity, with a mean August 1985 Crayfishes and Shrimps of Illinois 369 Fig. 96,—Distribution of Cambarellus puer in Illinois; black dots = 1972-1982 collections. growth increment of 0.45 mm 'molt (Black 1966). .Sixteen form I males col- lected in Illinois ranged from 7.7 to 12.8 and averaged 10.4 mm CL. In Louisiana, females carrying eggs or young occur January-April, and August-September (Penn 1950). In Illinois, females carrying eggs were collected Marcfi-May, and females carrying young have been collected in May (Table 6). Ten Illinois females carrying eggs (collected March-May) ranged from 11.0 to 14.6 mm and av- eraged 12.2 mm; five females carrying young (in May) ranged from 10.6 to 11.5, and averaged 10.9 mm CL. Six Illinois females, 11.0-14.6 mm CL, had 52-98 (mean, 82.3) eggs av- eraging 1.1 mm in diameter. Five fe- males, 10.6-11.5 mm CL, had 15-48 (mean, 30.8) young attached; the young on the female with 48 yrning were first instars, the 15-36 young on the other females were later instars, suggesting a high mortality while they are carried by the mother. Of the 60 C. puer collected in Illinois, 30 were males (16 were form I), and 30 were females. Longevity is 15-18 months (Black 1966). Genus Procambarus Ortmann Procambarus Ortmann 1905 Paracambarus Ortmann 1906 Orlmannicus Fowler 1912 Gonopod terminates in three or four (sometimes two in species ocdirring outside of Illinois) very short elements. Ischia of third, or third and fourth, pereiopods of male with hooks. Illinois harbors four species of Pro- cambarus, a genusof 148sije(ies(Hobbs 1981) reaching its greatest diversity in the southern United States and extend- ing as far south as Cluba and Honduras. The lour s|3e( ies in Illinois ari' sluggish- water inhaljitanis, as are most species of the genus. Illinois species are in the subgenera Girardiella (gracilis), Ort- niinuiicus (aculus and iiian'iridis). and .S( afj u lit a )n bams (da rk li). 370 Illinois Natural History Survey Bulletin Vol. 33. An. 4 Procambarus gracilis (Bundy) (Fig. 98) Catnbarus gracilis Bundy 1876 Description.—Rostrum broad, deejjly exca\aied. with short acumen, no median carina; margins barely converge anteriorly, without tubercles or spines. Carapace compressed, lacking cervical spines; suborbital margin angular. Areola extremely narrow, almost linear. Chela large, hea\ily punctate; palm with one row or large tubercles on mesial maigin; dactyl with shallow concavity on basal half of opposable margin. Form I gonopod terminates in four short elements: a sclerotized, curved central projection; a curved mesial process; a straight cephalic process; Fig. 97.— Bell Pond, east of Grantsburg, Johnson County, Illinois (8 October 1984), supports large populations of C. puer and P. clarkii. August 1985 Crayfishes and Shrimps of Illinois 371 Table 6.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Cambarellus puer. 372 Illinois Natural History Survey Bulletin \'ol. 33. An. 4 I Fig. 98 —Procambarus gracilis. A, dorsal view of carapace: B, dorsal view of right chela: C, annulus ventralis: D, mesial view of gonopod of form I male: E. lateral view of gonopod of form I male: F, lateral view of gonopod of form II male. August 1985 Crayfishes and Shrimps of Illinois 373 Fig. 99.—Total distribution of Procambarus gracilis. Ttie eastern limit of its range is un- known. Illinois in February, March, April, and May. Brown (1955) found form I males in Illinois in August and October and females with young attached in March, April, and October. Brown's largest form I male was 39 mm CL, and his largest female was 47 mm CL. Williams & Leonard (1952) found young attached to many of the females collected in early spring in Kansas. Greaser (1932) Fig, 100.— Distribution of Procambarus gracilis in Illinois. Open circle = 1922 record for the Kankakee River; black dots = 1972-1982 collections. noted that "females with young at- tached are taken in Missouri as late as Octobei." Among all specimens collected in the present study, males outnumbered Table 7.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Procambarus gracilis. 374 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 4 <-:.'k^' Fig. 101 .—Roadside ditch, % mi. W Winterrowd, Effingham County, Illinois. 3 f^/lay 1977; typi- cal habitat for Procambarus gracilis. Fig. 102.—Capped entrance to burrow of Procambarus gracilis in a field in Des Plaines, Cook County, Illinois, 15 June 1972. Males ( Form I) _i ca 5 -1 5 rH T Females if Vl.n - 1 nn„^^ 5 10 15 August 1985 Crayfishes and Shrimps of Illinois 375 Fig. 104. — Procambarus acutus. A, dorsal view of carapace: B, dorsal view of right chela; C. annulus ventralis; D, mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. 376 Illinois Natural History Survey Bulletin Vol. 33, An. 4 Procambarus acutus (Girard) (Fig. 104) Cambarus acutus Girard 1852 Cambarus stygius Bundy 1876 Description.—Rostrum broad pos- teriorly, strongly converges anteriorly, excavated, no median carina, marginal spines or tubercles small to absent (largest on small individuals). Carapace compressed, cervical spines present; suborbital margin angular. Areola narrow, narrowest part about 5-8 per- cent of length. Chela long and narrow, heavily piinctate; palm with row of large tubercles on mesial margin, one or two adjacent rows on dorsal and ventral surfaces. Form I gonopod terminates in four elements, all more or less directed caudodistally and vari- ously obscured by a mass of setae origi- nating on the caudal knob: a relatively large sclerotized central projection, an unsclerotized slender mesial process, a sclerotized cephalic process, and a sclerotized caudal process. Dorsal color pinkish brown to dark red; largest in- dividuals dark red with a black rectangle on each abdominal segment. Fig. 105.—Total distribution of Procambarus acutus. P. acutus is a member of the P. blan- dingii species group of the subgenus Ortmannicus (Hobbs 1962, 1972b). Two subspecies of P. acutus are recognized. P. a. cuevachicae occupies San Luis Potosi and Puebla and intergrades with P. a. acutus in northern Mexico and Texas. P. a. acutus occupies the rest of the range of the species (Hobbs 1972a), as described below. Hobbs ( 1981 ) noted the existence of se\eral regionally restricted "variants" of P. a. acutus^nd suggested that some may deserve taxo- nomic recognition. Cambarus stygius was described bv Bundv (in Forbes 1876) from specimens found, following a storm, on the shore of Lake Michigan at Racine, Wisconsin. (Bundv 1882) and listed by Forbes (18/6) as an Illinois species. Creaser (1932) declared Cambarus stygius to be a syno^ nym of Cambarus blandmgii acutus (- Cambarus acutus Girard). Distribution and Habitat.— P. acutus has a disjunct range. In the east it occurs along the Atlantic Slope from Maine to central Georgia (Altamaha Rner sys- tem) and to the west (Great Lakes and Mississippi Ri\er basins) it ranges from southern Michigan and western Indiana west to southeastern Minnesota and south to Alabama, western Oklahoma, and along the Gulf slope into Mexico (Fig. 105). P acutus has been counted among the Illinois fauna since 1870 (Hagen 1870), and Forbes (1876) stated that it was very common in ceniral Illinois. Rietz (1912) found the species (as Cam- barus blaminigii) to be widely distrib- uted and generally abundant and docu- mented its occurrence in ;}2 Illinois counties. Brown (1955) found it m 36 counties drained by the Sangamon Wabash, and Ohio river systems and reported it especiallv common in the Sangamon system. Populations m Il- linois were generally referred to as P blaniiingii until Hobbs (1962) re- stricted the range of P. a. blanclmgn) to North and South Carolina. Presently P. acutus occupies most, perhaps all, drainages of the state, but August 1985 Crayfishes and Shrimps of Illinois 377 Fig. 106 —Distribution of Procambarus acuius in Illinois; black dots = 1972-1982 col- lections. records are absent from the Spoon and other small rivers in western Illinois and from the Pecatonica and other streams in northwestern Illinois (Fig. 106). It is most common in the sluggish streams of southern Illinois, especially in the Big Muddy drainage and in streams on the Coastal Plain. Being a relatively deep water species, P. acutus is some- what more difficult to capture than are shallow water crayfishes, and it may be more common in Illinois than present records suggest. In Illinois, the species is most com- mon in permanent standing water bodies, especially those with luxuriant vegetation, and also is found in stream pools and slowly flowing runs with mud or sand bottoms. Elsewhere, the species occupies similar habitats but also may rarely be found in swiftly flowing streams (e.g., Williams 1954; Brown 1959; Hobbs 1981). A burrow may be constructed and inhabited when water bodies reach below-normal levels (Cralley 1932; Creaser &: Ortenburger 1933), adults may burrow during cold weather, and females sometimes sequester themselves in burrows when carrying eggs and young (Cralley 1932; Penn 1956; Hobbs 1981). The sequestering behavior could explain the extremely small number of ovigerous females found in Illinois and elsewhere (see below). In the Reel- foot Lake area, Hobbs & Maichand (1943) found burrows of P. acutus to be shallow excavations, usually under logs and consisting of a single passage- way extending only a few inches under- Table 8.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Procambarus acutus. 378 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 ground. A burrow 60 cm deep was found along the margin of a drainage ditcfi in Iowa (Phiillips 1980). Life History.—In the present survey of Illinois, 212 collections of P. acutus were made. Form I males were found every month except October and No- vember (Table 8). They peaked in rela- tive abundance in June-September and, secondarily, in January. Cralley (1932) found form I males in Illinois in June and July, and Brown (1955) from April to July and in September. Form I males have been found in June and July in western Tennessee (Hobbs &: Marchand 1943); in August and September in the Ozark-Quachita region (Williams 1954); from February to July and in September in Texas (Penn & Hobbs 1958); from May to August in Iowa (Phillips 1980); in April, August, and "the fall" in Georgia; in March, April, June, July, and September in Alabama; in April and May in Florida; and in all months except May in South Carolina (Hobbs 1981). No females were found carrying eggs, and only one (49.3 mm CL), collected in December, was carrying young {N = 30). In other Illinois surveys, Rietz (1912) reported a female collected with young in April, Cralley (1932) reported ovigerous females collected in July, and Brown (1955) found a female with eggs in August and one with young attached in April. All females with eggs and young found by Cralley and Brown were taken from burrows. Few females with eggs or young have been collected elsewhere, probably because of their sequestering behavior (Penn 1956; Hobbs 1981). In\he south- eastern United States, no females with eggs and only one female with young (from Alabama in September) have been collected (Hobbs 1981). Turner (1926) reported eggs and young on P. aculus in Ohio and Indiana in March, July, and September, and Penn (1956) found females with young attached in October and January in Louisiana. In the present survey males slightly outnumbered females, 523 to 513. The largest Illinois specimen is a 54.1-mm CL female collected on 20 February 1974 in the Maeystown Creek system, Monroe County. The largest male is a 54.0 mm-CL form I collected in 1974 in the Mississippi River, Whiteside County. Hobbs (1981) recorded a 58.9- mm CL form I male from Georgia. The smallest form I male from Illinois is 30.8 mm CL and was collected in the Mississippi River, Randolph County, on 24 March 1975. Procambarus viaeviridis (Faxon) (Fig. 107) Cambarus viae-vnidis Faxon 1914 Description.—Rostrum flat, deflect- ing downward anteriorly; margins con- verge anteriorly to short acumen, with- out spines or tubercles; no median carina. Carapace compressed, lacking cervical spines; suborbital margin barely angular. Areola narrow, narrow- est part about 4-7 percent of length. Chela long and narrow, heavily punc- tate; palm with row of 7-9 long tuber- cles on mesial margin. 2-3 adjacent rows of smaller tubercles. Form I gono- pod terminates in four short elements: a sclerotized mesial process, a sclero- tized central projection, a cephalic process, and a large setae-covered caudal knob on cephalic surface. Adults are deep rust red dorsally with the ab- domen somewhat lighter in color than the carapace. P. viaeviridis is a member of the P. blandingii species group of the sub- genus Ortmanniciis (Hobbs 1962, 1972b). Distribution and Habitat.—P. viae- viridis occupies standing water bodies and sluggish streams on the former Mississippi Embayment and Gulf Coastal Plain from southern Illinois south to northern Louisiana and cen- tral Alabama (Fig. 108). P. viaeviridis is restricted in Illinois to cypress swamps and floodplains along sluggish streams in the extreme southern part of the state (Fig. 109). First reported for Illinois by Page and Burr (1973), the species now is known August 1985 Crayfishes and Shrimps of Illinois 379 Fig. ^07.—Procambarus viaeviridis. A, dorsal view of carapace; B, dorsal view of right chela- C, annulus ventralis; D, mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. 380 Illinois Natural History Survey Bulletin 1 Vol. 33, An. 4 Fig. 108.—Total distribution of Procambarus viaeviridis. from localities in the Ohio River drain- age in Alexander, Johnson, Massac, Pope, and Pulaski counties. The largest populations are in cypress swamps {especially Heron Pond-Little Black Slough and Bell Pond) in Johnson County. Life History.—Other than brief comments on its affinity for sluggish and standing bodies of water, nothing appears to have been published on the ecological characteristics of P. inaeviri- dis. During the present study, 16 collec- tions of the species were made, all from December through May (Table 9). Fig. 109 —Distribution of Procambarus viaeviridis in Illinois: black dots = 1972-1982 collections. During this period, water levels in southern Illinois are generally high, and P. I'iaei'iridis can be found on flooded floodplains, in some roadside ditches, and in swamps. Periodically flooded areas appear to be the preferred habitat of P. I'iacviridis, and when they dry, (he crayfish burrows underground. In April, specimens were dug from bur- rows 15-30 cm deep on the floodplain Table 9.—Frequency of occurrence of form I males, females carrying eggs (ovigerous). and females carrying young, in Illinois collections of Procambarus viaeviridis. August 1985 Crayfishes and Shrimps of Illinois 381 '> ~' "^-^i i^rT'^^^ , i.-* •*<<.• \L.. V ^,H ^, '• S^i. Fig. 110.—Floodplain of Max Creek at the Route 147 bridge, Johnson County, Illinois, 8 October 1984. Procambarus viaeviridis can be found here above ground during flooding and can be dug from burrows during periods of low water. P. viaeviridis persists here even though the formerly forested floodplain now is covered with grasses. of Max Creek, Johnson County (Fig. 110). In December and January, 1975- 76, crayfish were active in water 1°C under a cover of ice. The usual absence of floodwater in southern Illinois from June through November accounts for the lack of Illinois collections during this period (Table 9); howe\er, more extensive effort presumably would Males ( = Form I) mm CL Fig. 111.—Size-frequency distribution of Procambarus viaeviridis collected in Bell Pond, Johnson County, Illinois, 19 May 1973. reveal that the species may be found in permanent swamps even during this period. Form I males were present in collec- tions made in January and May (Table 9). Among the four form I males col- lected, the smallest was 24.1 mm CX, and the largest was 29.9 mm C^L. I'he largest Illinois specimen of P. viaeviri- dis is a 32.3-mm CL female collected in Heron Pond on 30 March 1973. No females with eggs or young attached were collee ted. The smallest indi\ iihials (about .5-10 mm CL) were collected in Januaiy and P'ebruary. riie leiigth-iieciuency distiibution of indi\iduals (ollected in Bell Pond in May (N - 50) suggests that the spec ies, at least in Illinois, lives 2 years (Fig. 111). The sex ratio in this collection was 1 .1 males: 1 female. Procambaru.s ciarkii (C»iiard) (Fig. 112) Ciimhariis Clarkii Ciiard 1852 Description.— Rostrum dee|)ly ex- 382 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 Fig. ^^2.—Procambarus clarkii. A, dorsal view of carapace: B, dorsal view of right chela; C, annulus ventralis; D, mesial view of gonopod of form I male; E. lateral view of gonopod of form I male; F. lateral view of gonopod of form II male. August 1985 Crayfishes and Shrimps of Illinois 383 cavated; margins converge anteriorly, terminating in small to large spines; acumen long; no median carina. Cara- pace compressed, with cervical spines; suborbital margin angular. Areola ex- tremely narrow, sometimes obliterated at midlength. Chela long and narrow, punctate; palm with row of 6-8 tu- bercles on mesial margin, 2-3 adjacent rows of smaller tubercles. Form I gono- pod with a large angular shoulder on cephalic surface, a large lobiform scle- rotized cephalic process, small mesial process, and small sclerotized central projection. Light to dark red dorsally and laterally; white ventrally with blue stripe along middle of abdomen. P. clarkii is a member of the subgenus ScapuUcarnbarus (Hobbs 1972b), an assemblage of six species confined to the Atlantic and Gulf Coastal Plain of the United States and Mexico (Hobbs & Grubbs 1982). Distribution and Habitat.—P. clarkii is an abundant inhabitant of standing water bodies and slowly flowing streams on the Gulf Coastal Plain from the Florida panhandle to northern Mexico and up the former Mississippi Embay- ment to southern Illinois (Fig. 113). It is the crayfish commonly harvested for human consumption in the eastern United States. In Illinois, P. clarkii occurs in swamps, vegetated ponds, and stream pools in the extreme southern part of the state (Pope, Johnson, Massac, Union, Pulaski, and Alexander coun- ties) (Fig. 114). Most localities are in the Ohio River drainage, although some are in the Mississippi River drain- age. It remains common, especially in Mermel Swamp (Fig. 89), Bell Pond (Fig. 97), LaRue Swamp, Horseshoe Lake, and parts of the Cache River sys- tem but undoubtedly was more com- mon prior to the drainage of much of the southern Illinois wetlands. P. clarkii is taken often with P. acutus but is more common than P. acutus in the cypress- Fig. 113.—Total distribution of Procambarus clarkii. The southwestern limit of its range is unknown. Fig. 1 14.— Distribution of Procambarus clar- kii in Illinois; black dots = 1972-1982 collec- tions. 384 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 Table 10.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Procambarus clarkii. August 1985 Crayfishes and Shrimps of Illinois 385 Fig. 116. — Orconectes illinoiensis. A, dorsal view of carapace; B, dorsal view of right chela; C, annulus ventralis; D, mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. 386 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 found two females witfi eggs and three with young attached in Illinois on 6 September 1954; all were dug from bur- rows. Penn (1943) found a female only 30.5 mm CL carrying young in Loui- siana. The largest Illinois specimen is a 60.7-mm CL female collected in Horse- shoe Lake, Alexander County, on 20 June 1973; the largest male, from the same collection, is a 51.7 inm CL form I. Brown (1955) recorded a 54- mm CL form I male from Illinois. The size-frequency distribution of 116 individuals collected in the Bay Creek system on 19 May 1973 suggests that individuals of both sexes live a maxi- mum of about 2 years (Fig. 115), al- though the largest female mentioned above, at 60.7 mm, may have been 3 years old. Genus Orconectes Cope Orconectes Cope 1872 Faxonius Ortmann 1905 Gonopod with two terminal ele- ments; central projection not large and bladelike. Ischium of third pereio- pod with hook. Ten of the 65 recognized species of Orconectes (Hobbs 1981) occur in Il- linois. Species of Orconectes are, with few exceptions, open stream inhabitants and they, more than other crayfishes, have foimd Illinois suitable for oc- cupation. Orconectes illinoiensis Brown (Fig. 116) Orconectes illinoiensis Brown 1956 Description.—Rostrum deeply exca- vated, deflecting downward anterior- ly; long acumen; margins thick, con- verging anteriorly, straight to slightly concave, terminating in spines; no median carina. Carapace flattened dorsoventrally, with large cervical spines; suborbital margin smooth to slightly angular. Areola moderately wide, narrowest part about 13-15 per- cent of length. Chelae large, heavily punctate; fingers often very long; palm with row of 8-9 tubercles on mesial margin; 1-2 adjacent rows of tubercles on dorsal surface. Form I gonopod ter- minates in two short, straight elements: a sclerotized central projection and an unsclerotized mesial process. Dorsally light to dull red brown; fingers of chelae with red tips, subdistal black bands. O. illinoiensis is a member of the Propinquus group of Orconectes (Fitz- patrick 1967). The closest relative in Illinois is O. propinquus. Distribution and Habitat. — O. il- linoiensis is endemic to southern Il- linois. It was recognized first by Brown (1955) as a distinct species and formally described by him in 1956. During the present survey it was found to occupy rocky riffles and pools of streams in the Ohio and Big Muddy (Mississippi River drainage) systems of Hardin, Pope. Johnson, Massac, Pulaski, Alexander. Union, Williamson, and Jackson counties (Fig. 117). Most of the known Fig. 117— Distribution of Orconectes illi- noiensis: blacl< dots = 1972-1982 collections. August 1985 Crayfishes and Shrimps of Illinois 387 localities (Fig. 118) are in the Shawnee Hills, but a few populations occupy rocky streams on the Coastal Plain and in the region just north of the Shawnee Hills. Some populations may have been eliminated or reduced in recent decades through stream desiccation or other recent perturbations, but the distribu- tion of the species today probably ap- proximates its prehistoric distributicjn. Many populations are large, especially in upland streams in Pope County. Brown (1956) recorded a collection of O. illinoiensis from Cypress Ditch near Junction in Gallatin County. This is in the Saline River drainage, from Fig. 118 —Gibbons Creel*, Herod, Pope County, Illinois, (8 October 1984) provides habitat for Orconecles illinoiensis. 388 Illinois Nati'ral History Sirve^- Billetin \'f)l. 33, An. 4 Table 11.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Orconectes illinolensis. August 1985 Crayfishes and Shrimps of Illinois r Ji?V^°~?'?°"l'"®^ immunis. A, dorsal view of carapace: B. dorsal view of right chela- rormTma eT LtPr«, ""'f "'" °' ^°"°P°' °' '"^"^ ' "^^'^^ ^' '^'«-' -«- °' gonop'od ojlortn 1 male. F, lateral view of gonopod of form II male. 390 Illinois Natural History Survey Bulletin Vol. 33, An. 4 Fig. 121. immunis. -Total distribution of Orconectes spines; suborbital margin bluntly an- gular. Areola narrow, constricted an- teriorly, narrowest part about 10 percent of length. Chelae moderate in size, heavily punctate; palm with 2-3 rows of small tubercles on dorsomesial mar- gin. Form I gonopod terminates in two fairly short (central projection about 23 percent of total length of gonopod) elements curved at 90° angle to main axis: a sclerotized central projection and an unsclerotized mesial process. Annulus ventralis distinctive, with fos- sa far to one side. Dorsal color highly variable, from uniformly red brown to light green with dark brown mottling; fingers of chelae without bright red tips and subdistal black bands. O. immunis is a member of the J'irilis group of Orconectes and is most similar to O. alabamensis, O. rhoadesi, and O. validits. spec ies inhabiting the Caim- berland, Tennessee, and Mobile Bay drainages. Some populations of the wide-ranging O. immunis have been considered sufficiently distinct to war- rant laxonomic recognition (see syn- onymy abo\e), and Bovbjerg (1952) referred to northeastern Illinois popu- lations as "O. immunis x pedianus." However, Williams &: Leonard (1952) found that the characteristics which had been used to distinguish subspecies varied clinally and could not be used to diagnose geographically limited popu- lations. No subspecies were recognized by Hobbs (1974b). Distribution and Habitat. — O. im- munis ranges from southern Quebec, Maine, and Connecticut west through the Great Lakes (except Lake Superior), to eastern Wyoming and eastern Col- orado (Fig. 121). The southern edge of the range approximates the southern limit of Pleistocene glaciation but extends farther south into western Ken- tucky and Tennessee and farther west into the Great Plains. An isolated pop- ulation in Muddy Creek (Canadian- Arkansas River drainage), Okfuskee Fig. 122.— Distribution of Orconectes im- munis In Illinois; black dots = 1972-1982 col- lections. August 1985 Crayfishes and Shrimps of Illinois 391 % ^^ Fig. 123— Little Wabash River, 2 mi. WSW Mattoon, Coles County, Illinois, 25 October 1983. This and other sluggish weedy streams in Illinois often contain large populations of Orconectes immunis. County, Oklahoma, (Creaser &: Orten- burger 1933) is far south of other popu- lations and may have been introducted. The species occurs throughout Illi- nois except in the unglaciated north- west (Fig. 122). It appears to be rare in extreme western Illinois, although Rietz (1912) found it in several western counties (Fulton, Hancock, Henderson, and Mercer) for which recent records are unavailable. Elsewhere in the state it is generally common, and local popula- tions sometimes consist of thousands of individuals. It undoubtedly was even more common and widespread when Illinois had large expanses of undrained prairies. In fact, Forbes (1876) referred to O. immunis as "the commonest species of central Illinois," and "es- pecially frequent in the muddy ponds of the prairies, where it may be drawn by the hundreds with a small seine." O. immunis lives in slowly flowing pools, usually with soft bottoms, of headwaters, creeks, and srnall to large rivers (Fig. 123). It also is found in standing-water bodies but, at least in Illinois (and in the Reelfoot Lake region of Tennessee, Hobbs & Marchand 1943), is primarily a stream species. It fre- quents intermittent streams and tem- porary ponds, burrowing down to the water table as drying occurs. O. im- munis can be found under rocks in dry- ing streams, and burrows are con- structed as a means of escaping des- iccation. Burrows usually are simple structures, although they may have chimneys and occasionally are relatively elaborate. Brown (1955) described the burrows as up to 2 feet deep and ending in an enlarged pocket below the water table. Tack ( 1 94 1 ) made observations on the 392 Illinois Natural History Sirvev Bi lletin \ol. 33. An. i Table 12.—Frequency of occurrence of form I males, females carrying eggs (ovigerous). and females carrying young, in Illinois collections of Orconectes immunis. August 1985 Crayfishes and Shrimps of Illinois Fig. 124.— Orconecres indianensis. A, dorsal view of carapace; B, dorsal view of rigfit cfiela; C, annulus ventralis; D, mesial view of gonopod of form I male: E. lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. 394 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 total length (about 20 mm CL) (Rietz 1912). Females carry eggs in Illinois from January through April and carry young from April through June (Rietz 1912; Brown 1955; Table 12). The smallest ovigerous Illinois female is 22 mm CL (collected in Richland County, 12 April 1979), the same length as that reported by Tack (1941) in New York. Elsewhere, eggs have been noted in April and November in Michigan (Her- rick 1896; Pearse 1910); April, May, and October in Iowa (Caldwell & Bovb- jerg 1969; Phillips 1980); March and April in Indiana (Williamson 1907); April in Kansas (Harris 1902); April and October in Nebraska (Engle 1926); June in North Dakota (Creaser &: Orten- burger 1933); and October-November and April-May in New York (Tack 1941; Crocker 1957). Young are carried in May in New York and Ontario (Tack 1941; Crocker 1957; Crocker &Barr 1968) and from April through June in Iowa (Caldwell Xc Bovbjeig 1969; Phillips 1980). Seven Illinois females collected dur- ing the present study (21.8-35.9 mm CL) were carrying 102-285 (mean, 168.9)eggs. Eggs averaged about 1.7mm in diameter. Three females (26.6-35.5 mm CL) carried 58-184 (mean, 120.3) young. The largest individual encountered during the present survey was a 41.9- mm CL form II male collected in Ben- nett Creek, Crawford County, on 30 April 1973. Brown (1955) collected a 46-mm CL female from Illinois, and Crocker (1957) recorded a 49-inm CL female from New York. The ratio among all Illinois specimens collected was 505 males: 513 females. Orconectes indianensis (Hay) (Fig. 124) Cainbarns itidianensis Hay 1896 Description.— Rostrum exca\ated, slightly deflet ted downward anteriorly, with long acumen: margins con\erge slightly anteriorly, ending in small Fig. 125.—Total distribution of Orconectes indianensis. spines; small median carina. Carapace flattened dorsoventrally. with large cervical spines: suborbital margin barely angular. Areola moderately wide, narrowest part 15-20 percent of length. Chela large, hea\ily punctate; palm with row of 7-9 tubercles on mesial margin. 2-3 adjacent rows of smaller tubercles on dorsal surface. Form I gonopod terminates in two short, distally tapering and strongly di\erging elements: a sclerotized cen- tral projection and an imsclerotized mesial process. Overall dark brown dorsally, except thorax yellow brown; fingers of chelae with red tips and sub- distal black bands. O. indianensis is most similar, and perhaps most closely related, to O. wrighli. a species known only from Hardin Countv, Tennessee (Hobbs 1974b). Distribution and Habitat.—O. in- dianensis lives in rocky riffles and pools of small to medium-sized streams in the Wabash Ri\er drainage of southwestern Indiana and in the .Saline Ri\er and Honey Creek systems (both tributary to August 1985 Crayfishes and Shrimps of Illinois 395 the Ohio River) in southeastern Illinois (Fig. 125). This species uses stones as cover and often constructs a short tunnel with a terminal pocket under a stone as a resting place (Brown 1955). Recent Illinois collections of O. in- dianensis are available from the Saline drainage of Gallatin, Saline, William- son, Pope, and Johnson counties, and from Honey Creek in Hardin County (Fig. 126). Most collections in the Saline system are from that part of the system draining the Shawnee Hills. The north- ern part of the system, draining the Mount Vernon Hill Country (Central Lowland Province) seems to be devoid of O. iudtanensis. Honey Creek, also in the Shawnee Hills, is the first large stream south of the Saline River. Brown (1955) documented essentially the same distribution in Illinois as did the present survey, but his record for O. illinoiensis in Cypress Ditch, near Junction, Gallatin County was prob- ably based on O. indianensis. Rietz (1912) had more widespread records, but some of them probably (because of the localities) were based on misidentifica- tions of the then undescribed O. illi- noiensis and O. stanndardi. Rietz's record for Grand Pierre Creek, Pope County, and two records for Bay Creek, Pope and Johnson counties were surely based on O. illinoiensis. Her record for Skillet Fork, Wayne County, was probably based on O. stannardi. The Little Fox River, White County, record may have been based on O. propinquus. However, her records for South Fork Fig. 126.—Distribution of Orconectes indi- anensis in Illinois. Open circles = pre-1912 records (Rietz 1912); black dots = 1972-1982 collections. Saline River and, more interestingly, for the North Fork Saline River must have been for O. indianensis. No simi- lar species occurs or ever has been re- corded for the Saline system. The North Fork records are interesting because O. indianensis no longer is found there. The North and Middle forks of the Table 13.—Frequency of occurrence of form I males, females carrying eggs (ovigerous). and females carrying young, in Illinois collections of Orconectes indianensis. 396 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 I^ Males Form I) „ [0 Femal es ( with eggs) 10 20 30 mm CL 40 Fig. 127.—Size-frequency distribution of Orconectes indianensis collected in Honey Creek, Hardin County, on 23 Marcfi 1973. .Saline are badly polluted. They suffer from strip-mine and oil-field pollution, siltation, and desiccation (Smith 1971) and are among the most damaged and abused streams in Illinois. In many areas of these streams little aquatic life of any kind can be found. Life History.—Apparently the only ecological information on O. indianen- sis is that presented by Brown (1955) and that from the present survey. Brown (1955) found no ovigerous females and collected form I males only in September and November. In the present study, collections were made from March through October (except June), and form I males were present in March, August, September, and October (Table 13). The smallest form I male was 17.7 mm CL, and the largest was 33.4 mm CL. Ovigerous females were found in March and April (Table 13). One hun- dred twenty-one eggs were counted on a 26.3-mm CL female, 148 eggs on a 27.7-mm CL female, and 178 eggs on a 32.2-mm CL female. Eggs averaged about 1.8 mm in diameter. The only female with young attached was 35.0 mm CL and was collected on 19 May 1973. Attached to her were 132 young and 21 unhatched eggs. The largest Illinois specimens ex- amined were the above-mentioned 35.0- mm female and 33.4-mm form I male. The size-frequency distribution of 51 specimens collected in Honey Creek. Hardin County, on 23 March 1973 (Fig. 127) suggests a 2-year life span, with 1- year-old males averaging 15.2 mm (N = 25), 1 -year-old females averaging 13.4mm(N = 22),and2-year-olds(.V = 4) ranging from 25 to 33 mm CL. Of the 219 specimens of O. indianensis col- lected during the present survey, 104 were males and 115 were females. Orconectes kentuckiensis Rhoades (Fig. 128) Orconectes kentuckiensis Rhoades 1944 Description.—Rostrum excavated, deflected downward anteriorly, with long acumen; margins barely converge anteriorly, ending in tubercles or small spines; lacks median carina. Carapace flattened dorsoventrally, with large cervical spines; suborbital margin barely angular. Areola wide, narrowest part about 18 percent of length. Chela large, hea\ily punctate; palm with row of 7-9 tubercles on mesial margin; 1-2 adjacent rows of smaller tubercles on dorsal surface. Form I gonopod termi- nates in a short, thick, distally directed, sclerotized central projection, and a short, caudodistally directed, unsclero- tized mesial process. Dorsally dark brown (with yellow brown thorax) to red brown; fingers of chelae with red tips, subdistal black bands. The gonopod of O. kentuckiensis most closely resembles that of O. sloanii in southern Indiana and Ohio and O. hamsonii in southeastern Missouri. Although the relationships of these species ha\e not been in\estigated. O. sloanii and O. harrisonii probably are the closest relatives of O. kentuck- iensis. Distribution and Habitat. — O. ken- tuckiensis occurs only in small streams in a small area of southeastern Illinois and northwestern (Crittenden. Li\ing- ston, and I'nion counties) Kentucky (Fig. 129). In Illinois, it occurs only in Big, Hosick, and Peters creeks, three spring-fed, rocky, direct tributaries of the Ohio River in Hardin County (Fig. 130). Its distribution in the headwaters of Big Creek (Fig. 131 ). by far the largest of the three streams, complements the downstream distributit)n of O. plandus I August 1985 Crayfishes and Shrimps of Illinois 397 Fig^ ^28.-Orconectes kentuckiensis. A, dorsal view of carapace: B, dorsal view of rioht chela; C annulus ventralls; D. mesial view of gonopod of form I male: E, lateral view of gonopod ot form I male, F, lateral view of gonopod of form II male. 398 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 Fig. 129.—Total distribution of Orconectes kentuckiensis. (Boyd & Page 1978). No record of its occurrence in Illinois predates those of Brown (1955). O. kentuckiensis prob- ably was never more widespread in Il- linois than it is today. O. kentuckiensis is most common in shallow, rocky pools of small streams (Boyd & Page 1978), where large rocks are used as cover, although Rhoades (1944a) collected specimens in accumu- Fig. 130.— Distribution of Orconectes ken- tuckiensis in Illinois; black dots = 1972-1982 collections. lations of brush over mud. This species does not burrow but may bury itself in loose gravel 2-4 cm below the surface of a dry stream bed (Boyd k Page 1978). Life History.—The life history of O. i { Table 14.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Orconectes kentuckiensis. Fig. 131—Headwaters of Big Creek, 2 mi E Karbers Ridge, Hardin County, Illinois, support large populations of Orconectes kentuckiensis. 400 Illinois Natural History Survey Bulletin \'ol. 33, Art. 4 kentuckiensis in Big Creek was studied by Boyd & Page (1978). The species was active at temperatures of 6°-32°C and dissolved oxygen levels of 3.5-14.4 ppm. Most individuals were found in water 5-61 cm deep. The species lived a maximum of 2+ years, had a 1:1 sex ratio, fed on vascular plants and arth- ropods (amphipods, isopods, crayfishes, caddisflies, and midges), and was preyed upon by fishes. Form I males (15.9-37.5 mm CL) were present from July through April and peaked in occurrence in November, December, and February; females carry- ing eggs or young were present in March, April, May, and July (Table 14) and measured 14.8-32.8 mm CL. Fif- teen females were carrying 49-248 (mean, 110) eggs. Eggs were spherical and averaged 1.9 mm in diameter. The relationship between the number of eggs (N) and the female's weight in grams (W) was N = 49.809 + 25.310 VV, r = 0.807, and that between N and the female's carapace length (L) was N = -0.43611 + 1.8516 log L, r = 0.859. Counts of ovarian eggs in 15 females were 59-249 and averaged 109. Nimi- bers of eggs, ovarian and abdominal, increased significantly with carapace length. The growth of first-year cray- fish occurred during a period of 4.5 months, of adult males during spring and summer molts, and of adult females during a summer molt. The largest O. kentuckiensis from Illinois is a 37.5-mm CL form I male collected on 25 April 1975; the largest female is 36.2 mm CL, collected on 30 September 1975. Both are from Big Creek. Orconectes lancifer (Hagen) (Fig. 132) Cambarus lancifer Hagen 1870 Cambariis faxonii Meek 1894 Description.—Rostrum deeply ex- cavated; acumen longer than basal mar- gin of rostrum, turning upward ante- riorly; no median carina; margins con- vex, diverging anteriorly, terminating in spines. Carapace slightly flattened dorsoventrally, with large cervical spines; suborbital margin angular. Areola narrow, obliterated anteriorly. Chela long and narrow, setose, without tubercles. Form I gonopod terminates in two short, spatulate elements: a sclerotized central projection and an unsclerotized mesial process. Mottled overall in shades of red, brown, and green; fingers of chelae without bright red tips and without black bands sub- distally. O. lancifer is a highly distinctive crayfish with no obviously close rela- tives. Distribution and Habitat.—O. lan- cifer is an inhabitant of the Gulf Coastal Plain and is known to occur in Louisi- ana, Mississippi, Tennessee, eastern Texas, Arkansas, and extreme southern Illinois (Fig. 133). It lives in deep water of oxbows, bayous, and large streams (Black 1972) and, as a result is rather poorly represented in collections. O. lancifer has been found at only two localities in Illinois. Robert Kenni- cott, an early naturalist, collected a female at Cairo in the middle 1800's (Faxon 1914). and Page & Burr (1973) recorded a collection of eight individ- uals from Horseshoe Lake. Both local- ities are in the Mississippi River diain- age of Alexander County (Fig. 134). Four additional collections of O. lan- cifer recently have been made at Horse- shoe Lake, where the species probab- ly is common but difficult to collect. Horseshoe Lake (Fig. 135) is an old. relatively deep oxbow of the Missis- sippi River supporting stands of bald cypress and man> species of Coastal Plain animals that are rare in Illinois (E\ers &: Page 1977). Few oxbows of the Mississippi River exist in southern Illinois, and O. lancifer is unlikely to be found elsewhere in the state. Wolf Lake in Lnion County appears to offer suitable habitat for O. lancifer. but several searches there ha\e failed to document its presence. In Louisiana. O. /fl;« //cr li\es in deep (more than 40 cm), clear, permanent August 1985 Crayfishes and Shrimps of Illinois r J'9, ^32.-Orconecfes lancifer. A, dorsal view of carapace; B. dorsal view of riaht chela- ?'lTl'itT\Z':;,°'. ---' -«- °' 9-opod of form I male; E. lateral v": o gonopod oflateral view of gonopod of form II male. form I male; F, 402 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 Fig. 133.—Total distribution of Orconectes lancifer. Fig. 134.— Distribution of Orconectes lanci- fer in Illinois. Open circle = collection made in the mid-1800's (Faxon 1914); black dots = 1972- 1982 collections. */:. W^^ -»% -^ Fig. 135 —Horsestioe Lake in Alexander County is the only known locality In Illinois still supporting a population of Orconectes lancifer. August 1985 Crayfishes and Shrimps of Illinois 403 Fig. 136. — Orconectes placidus. A, dorsal view of carapace; B C, annulus ventralis; D, mesial view of gonopod of form I male; E form I male; F, lateral view of gonopod of form II male. dorsal view of right chela; lateral view of gonopod of 404 Illinois Natural History Slirvey Bulletin Vol.33. Art. 4 water over mud or mixed mud and sand with little aquatic vegetation present (Penn 1952). This description fits Horseshoe Lake fairly well except that the lake is heavily vegetated. Life History.—Black (1972) sum- marized the scant life history data avail- able on O. lancifer: form I males have been collected from August through November; females carrying eggs or young have been found only in Feb- ruary; juveniles predominate in May, June, and July. Among the collections made in Il- linois are two made in June and one each in July, September, and October. The September and October collections contain a total of three form I males (30-32 mm CL). Also in the Illinois Natural History Survey collection is a 40-mm female from Cheniere Lake, Ouachita Parish, Louisiana, collected on 18 February 1977, with 570 eggs attached. The eggs average 1.5 mm in diameter. These additional observations agree with Black's (1972) description of an autumnal breeding season and winter egg-laying season. Orconectes placidus (Hagen) (Fig. 136) Cainbarus placidus Hagen 1870 Description.—Rostrum excavated, no (or low) median carina, long acu- men; margins thick, straight to slightly concave, barely converging anteriorly, terminating in spines. Carapace flat- tened dorsoventrally, with large cervical spines; suborbital margin smooth. Areola moderate, narrowest part about 11-15 percent of length. Chelae large, heavily punctate; palm with 2-3 rows of tubercles along mesial margin. Form I gonopod terminates in a long sdero- tized central projection curved at tip and a slightly shorter unsclerotizcd mesial process; cephalic surface smooth or with broadly rounded (not angular) shoul- der. Dorsal ly red brown; fingers of chelae with red tips, subdistal black bands. Fig. 137—Total distribution of Orconectes placidus. Fig. 138.— Distribution of Orconectes placi- dus in Illinois; black dots = 1972-1982 col- lections. August 1985 Crayfishes and Shrimps of Illinois 405 Fig. 139.— Gravel riffles in the lower portion of Big Creel<, Hardin County, Illinois (10 March 1976) provide the best habitat in Illinois for Orconectes placidus. Among the crayfishes of Illinois, populations herein assigned to O. placidus arc the most troublesome tax- onomically. The only large population is that in Big Creek in Hardin County; other populations are small, highly localized ones inhabiting rocky areas along the banks of the Ohio and Missis- sippi rivers. Although all are assigned to O. placidus, these populations show variations usually indicative of specific distinctiveness in Orconectes. The gonopod of the form I male lacks any hint of a shoulder on its dorsal surface in Big Creek specimens but has a defi- nite shoulder (although not as pro- nounced and angular as in O. rusticus) in Mississipi^i Ri\er specimens (no form I males from the Ohio River are avail- able). Big Creek specimens always lack a carina on the rostrum; those from the Ohio and Mississippi rivers may or may not have one. In general, western j)op- ulations differ the most from typical O. placidus occupying the Cumberland and Tennessee river systems in Ken- tucky and Tennessee (Hobbs 1971b). Variation in O. placidus and its rela- tives is badly in need of study. Distribution and Habitat.— O. placi- dus occupies streams of the Cumber- land, Tennessee, and lower Ohio rivers in Kentucky, Tennessee, and Illinois (Fig. 137). It lives in rocky riffles and pools, using cavities and interstices as hiding places. Recent Illinois records exist for Big Creek, Hardin County, the Ohio River, Massac and Pulaski counties, and the Mississippi River, Randolph and Jack- son counties (Fig. 138). Most of the large-river collections were made along rocky banks or in rocky backwater areas. In the Big Creek system, O. placidus is found mainly in downstream gravel and rubble riffles (Fig. 139), and O. kentuckiensis is found mainly in up- stream shallow rocky pools (Boyd &: Page 1978). As discussed below for O. rusticus, Hagen's (1870) and P'orbes' (1876) reference to "C. placidus" at Quincy is thought to refer to a waif of the Missouri po{)ulalions of O. ( f . rusti- cus. Brown's (1955) discussion of "O. 406 Illinois Natural History Survey Bulletin Vol. 33, An. 4 Table 15.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Orconectes placidus. Numbfr and Percent of Colleclions with Month Number of Collections Fovm I Males Ovigerous Females Females with Young I January February March April May June July August September October November December 3 1 6 2 6 19 3 1 7 8 4 1 67 83 71 88 75 100 17 50 <: a 1 r^MJu :;: es ( = Form I) 5 10 " 5 o ^ Females 10 1 ^ . n n 20 30 mm CL 40 Fig. 140.—Size-frequency distribution of Orconectes placidus collected in October in Big Creek, Hardin County. rusticus" refers to the Big Creek popu- lation of O. placidus. Life History.—During the life- history study of O. kentuckiensis {Boyd & Page 1978), a large number (61) of collections of O. placidus were made. The following observations are based on those collections. Form I males were present in collec- tions from September through January and in March (only one collection was made in February) (Table 15). Brown (1955) found form I males in Big Creek in November. Ovigerous females (N = 'i) were found only in March and April (Table 15); they were 15.7, 20.5, and 27.2 mm CL, with 12, 88, and 134 eggs attached, respectively. Eggs were spherical and averaged 2.1 mm in diameter. No fe- males with young attached were found. The largest pieser\ed Illinois male is a 45.6-mm form II; the largest female is 41.7 mm CL. Both were collected in May and, compared with the size-fre- quency distribution of October-col- lected specimens (the largest sample), were probably 2 or 3 yearsold(Fig. 140). The largest form I male (39.6 mm) has extremely large chelae (right chela = 4(>. 1 mm long). Among the Big Creek specimens were 344 males and 387 fe- males (a ratio of 1:1.13). Among a combined total of 506 stripe- tail and spottail darters {Etheosloma kennicotti and E. squainiceps) exam- ined, the ingested remains of crayfishes were found in the stomachs of 4 E. squainiceps (Page 1974a, 1975). Con- sidering the relative abundances of cray- fishes in Big Creek, these were almost certainly O. placidus or O. kentuckien- sis (Boyd & Page 1978). Orconectes propinquus (Girard) (Fig. 141) Carnbarus pwpitiquus Girard 1852 Orconectes iowaensis Fitzpatrick 1968 Description.— Rostrum exca\ atcd. deflecting downward anteriorly; well- developed median carina; long acumen; margins thick, straight or slightly con- tavc, barely coinerging anteriorly, ter- minating in spines or lagc tubercles. Carapace flattened doisoventrally, with August 1985 Crayfishes and Shrimps of Illinois 407 Fig. 141.—Orconecfes propinquus. A, dorsal view of carapace; B, dorsal view of right chela: C, annulus ventralis; D, mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. 408 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 cervical spines; suborbital margin smooth. Areola wide, narrowest part about 19-25 percent of length. Chelae large, heavily punctate; palm with 2-3 rows of tubercles on mesial margin. Form I gonopod terminates in two short, straight elements: a sclerotized central projection and an unsclerotized mesial process. Dorsal ly green to red brown; fingers of chelae with red tips, subdistal black bands. Fitzpatrick (1967) found O. propin- cjiius to be the most variable species among the Propinquus group of spe- cies but also to lack taxonomically recognizable subspecies. "O. wmaen- sis," previously thought to be a para- patric relative of O. propinquus oc- cupying areas of Iowa adjacent to Il- linois (Fitzpatrick 1968), has been rele- gated to the synonymy of O. propin- quus by Page (1985). Distribution and Habitat.—O. pro- pinquus ranges from southern Ontario and Quebec south to Illinois, Indiana, Ohio, northern Pennsylvania, New York, and Massachusetts. To the west, its range extends into eastern Iowa and southeastern Minnesota (Fig. 142). In Illinois, O. propinquus occurs in Lake Michigan and in the Wabash. Il- linois, Rock, Mississippi, and extreme upper Kaskaskia drainages (Fig. 143). In the Wabash drainage, it extends as far south as White County, is wide- spread in the Embarras River, but is absent in the Little Wabash River sys- tem. In the Illinois and Mississippi rivers, it occurs as far south as Calhoun and Rock Island counties, respectively. Throughout its range in Illinois, O. propinquus is the most common crayfish in clean rocky riffles (Fig. 144). In the fast, rubble and gravel riffles characteristic of the Vermilion, Kan- kakee, and upjH'r Rock drainagis, O. propinquus often is abundant. Its ab- sence in most of western and central Illinois presumably is due to the ab- sence of suitable habitat; its absence in southern Illinois apparently is due lo the presence there of t)lher species of Fig. 142.—Total distribution of Orconectes propinquus. The northeastern limit of its range is unknown. Fig. 143.— Distribution of Orconectes pro- pinquus in Illinois: black dots = 1972-1982. August 1985 Crayfishes and Shrimps of Illinois 409 Fig. 144.—Middle Fork Vermilion River, near Collison, Vermilion County, Illinois, provides abundant habitat for Orconectes propinquus. rocky stream-inhabiting species of Orconectes. O. proptuquus may dig under stones on the stream bed to escape desiccation (Bovbjerg 1952; Brown 1955) or occa- sionally into stream banks (Hay 1896; Cralley 1932). Young often are found in vegetation along the margins of a stream. Altliough primarily an inhabi- tant of rocky streams, it can become established in lakes (Bovbjerg 1952). O. propinquus remains u'idespiead and locally common, even abundant, in some areas but probably was more widespread and common prior lo the extensive modification of the Illinois landscape. It does best in clean, clear streams and presumably has suffered population declines as stream siltation has progressed during tfie last few dec- ades. Among the widespread crayfishes of Illinois, it is probably the best in- dicator of environmentally healthy streams. All earlier records of O. propinquus in Illinois fall within, or almost within, the present distribution. The record lor the Kaskaskia drainage, near Hum- boldt, Coles County (Rietz 1912), is one county to the south of recent records. This record is signifitani in document- ing tlie fact tliat the presence ol the 410 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 Table 16.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Orconectes propinquus. August 1985 Cravfishk and Shrimps of Iixih 411 Fig. 145—Orconecfes rusticus. A, dorsal view of carapace; B, dorsal view of right chela; C. annulus ventralis; D. mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. 412 Illinois Natural History Survey Bulletin Vol. 33, An. 4 and May in Iowa (as O. iowaensis, Phillips 1980), and eggs and young in April-June in Michigan (Pearse 1910; Creaser 1931). Females with eggs and young may be as small as 16 mm CL (Crocker 1957; Berrill 1978). Eggs are spherical and average about 1.9 mm in diameter. The ratio of males to females among recent Illinois collections is 334:335. The largest specimen collected during the present survey is a 39.3-mm CL female; the largest male is a 31.5-mm form I. Van Deventer (1937) and Brown (1955) each recorded a 40-mm male; Cralley (1932) recorded a 90-mm TL (about 45-mm CL) male and a 90-mm TL female from Champaign County. O. propinquus is omnivorous, feed- ing on plant materials and insect larvae (Bovbjerg 1952). Orconectes rusticus (Girard) (Fig. 145) Cambarus rusticus Girard 1852 Cambarus juvenilis Hagen 1870 Description.—Rostrum excavated; no median carina; long acumen; margins thick, concave, terminating in spines. Carapace flattened dorsoventrally, with cervical spines; suborbital margin smooth. Areola moderate, narrowest part about 10-15 percent of length. Chelae large, heavily punctate; palm with two rows of tubercles on mesial margin. Form I gonopod with an an- gular (90° to axis of gonopod) shoulder on dorsal surface, terminating in two long elements: a sclerotized central projection curved at its tip, and a slight- ly shorter unsclerotized mesial process. Dorsally green brown to rust red; often a large red spot surrounded by light brown on side of carapace; fingers of chelae with red tips, subdistal black bands. Close relationships to O. rusticus are not readily apparent. Species often confused with, and at times considered related to, O. rusticus include O. for- ceps, O. placidus, O. putnami, O. spinosus, and until synonymized with Fig. 146.—Presumed native distribution of Orconectes rusticus. O. rusticus by Bouchard (1977), O. juvenilis. Distribution and Habitat.— O. rusti- cus now occurs in New England. On- tario, Michigan, Ohio, Indiana, Ken- tucky, Tennessee, Illinois, Wisconsin, Minnesota, Iowa, Missouri, and New Mexico. It has been widely introduced, presumably through use as fishing bait, and the boundaries of its native range (Fig. 146) now are difficult to determine. However, at least its occurrences in New England (Crocker 1979), Ontario (Crocker & Barr 1968). Wisconsin (Capelli & Munjal 1982). New Mexico (Bouchard 1977). and Illinois are cer- tainly the results of introductions. Once introduced, O. ru5t?fU5 successfully dis- places native species and expands its range, apparently because it is a large and aggressi\e species able to thrive in impoundments and other disturbed habitats (Berrill 1978; Capelli 1982; Capelli & Munjal 1982: Capelli & Magnuson 1983). Its continued use as a li\e bait species is therefore unwise. Taxonomic confusion about O. rus- August 1985 Crayfishes and Shrimps of Illinois 413 Fig. 147.—Distribution of Orconectes rusti- cus in Illinois. Open circle = Adams County locality given by Hagen (1870) for "Cambarus placidus," black dots = 1972-1984 collections. ticus and its relatives has reigned since their descriptions. "Cambarus wiscon- sinensis Bundy," found at Normal (Forbes 1876), is a synonym of O. virilis (Hobbs 1972a) and is not a reference to O. rusticus, as thought by Rietz (1912) and Page (1974b). Brown's (1955) records for "O. rusticus" all were for Big Creek in Hardin County and are treated herein as O. placidus. "Cambarus placidus Hagen" was recorded from Quincy by Hagen (1870) and Forbes (1876) and considered by Rietz (1912) to represent a record of O. rusticus. Quincy is directly across the Mississippi River from the Fabius River system of Missouri which sup- ports a population of O. rusticus or a closely related undescribed species. The Quincy record probably represents a waif from the Fabius River population of Orconectes and is plotted on the O. rusticus map (Fig. 147). In Illinois, O. rusticushas been found recently in Peoria Lake, Tazewell County (in 1973 and 1974), Powder Horn Lake, Cook County (1975), Wolf Lake, Cook County (1975, 1978, 1980), Eagle Creek, Lake County (1975), Kent Creek, Winnebago County (1977), Nippersink Creek, McHenry County (1979), in a backwater of Fox River at Oak Point State Park, Lake County (1982), and in Lake Michigan, 1 mile east of Burnham Harbor, Cook County (1984). All of these localities are heavily used by fishermen or are near such areas, and the presence of O. rusticus in Il- linois is surely the result of bait-bucket introductions. Except for Lake Michi- gan and Wolf and Powder Horn lakes, which are in the Lake Michigan drain- age, and Kent Creek, which is a tributary of the Rock River, all localities are in the Fox-Illinois River system (Fig. 147). O. rusticus is reproducing in Illinois (at least in Wolf Lake, where specimens as small as 16 mm CL have been found) and probably is supplanting native species, as it is in southern Ontario (Berrill 1978) and Wisconsin (Capelli & Munjal 1982). Hybridization between O. rusticus and O. propinquus and between O. rusticus and O. limosus has been recorded in Wisconsin and Massachusetts (Capelli & Capelli 1980; Smith 1981) and appears to represent an intermediate step in the displace- ment by O. rusticus of these native species. In Illinois, it appears to be hybridizing with O. virilis in Wolf Lake. O rusticus lives in rocky riffles, pools, and along wave-swept lake margins. Life History.—In fish ponds in Ohio, Langlois (1935) found mating to occur in September and October; following copulation, the females burrow into banks. Oviposition occurs in a few females in October, but mostly in April and May, when females are 1 year old. Eggs hatch in about 20 days and remain attached to the female for al)out 5 days. Sexual maturity is reached in the fall 414 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 4 I Fig. 148— Orconecfes stannardi. A, dorsal view of carapace: B, dorsal view of right chela; C, annulus ventralis; D, mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. August 1985 Crayfishes and Shrimps of Illinois 115 following one summer's growth. Prins (1968) studied O. rusticus in a stream in Kentucky and found that mating peaks in September and Octo- ber, females oviposit from February through June at about 22-24 months, eggs hatch in May after being carried by the female for 4-6 weeks, young remain attached to the mother for 3 weeks {through April and May), sexual maturity is reached at about 15 months (about 18 mm CL), the sex ratio among juveniles is 1:1, and the species reaches a maximum longevity of about 3 years and a maximum CL of 45 mm. Counts of ovarian eggs in 43 females ranged from 54 to 357. The smallest female with eggs attached (55 eggs) was 17 mm CL; _the largest, with 231 eggs, was 38 mm CL. The relationship between the number of eggs (N) and carapace length (L) of 49 females was N = -91.49 + 8.3 IL, with r = 0.86. The species feeds mainly on plants and detritus and has been observed feeding out of water along the banks. During winter, O. rusticus typically burrows into stream banks and into interstices in the substrate. Form I males have been found in July and August in Ontario (Crocker Sc Barr 1968), in June in Maine (Crocker 1979), and in July and September in Iowa (Phillips 1980). Females in berry were collected in April in Iowa (Phillips 1980). Busch (1940) found that it took 20 Jays for eggs to hatch at 20°-22°C. .\mong the nine Illinois collections (made from May through September), form I males (20.5-34.0 mm CL) are present in August and September. No ovigerous females were found. Some Illinois specimens approach the maxi- mum size recorded for the species; the largest Illinois male (form II) is 40.5 mm, CL. and the largest female is 41.5 mm PageOrconectes stannardi (Fig. 148) Orconectes stannardi Page 1985 Description.— R(jstrum excavated; well-developed median carina; long acumen; margins moderately thick. subparallel, slightly converging me- dially, terminating in spines. Carapace flattened dorsoventrally, with cervical spines; suborbital margin weakly an- gular. Areola moderately wide, narrow- est part about 15-25 percent of length. Chela large, heavily punctate; palm with two rows of tubercles on mesial margin. Form I gonopod terminates in two short, straight elements: a sclero- tized central projection and an unsclero- tized mesial process with spur about middle of caudal surface; shoulder on cephalic surface at base of central pro- jection. Olive green to light brown and usually heavily speckled with dark brown dorsally; fingers of chelae with red tips, subdistal black bands. Distribution and Habitat.—O. stan- nardi is a member of the O. propincjuus species group that is endemic, and Fig. 149— Distribution of Orconectes stan- nardi. Open circles = localities given by Rietz (1912) and Brown (1955) for "O. propinquus" and "O. indianensis." black dots = 1972-1984 collections. 416 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 i^^f^^^W^^:^'''€l^^ '^aw 1 * „-w*k.: Fig. 150—Orconecfes stannardi is most common along rocky stream margins such as this one on the Little Wabash River, 3.5 mi NNW Louisville. Clay County, Illinois (8 October 1984). po.ssibly autochthonous, to the Little suggesting that the species oiiginated Wabash River system of Illinois. The sonietiine within the past 100.000 veais region drained by the Little Wabash (Page 198.")). .\11 collections made since was glaciated during the Illinoian 1972 are from the Little Wabash River (about 100,000 y.b.p.) but not the Wis- proper (Fig. 149). consinan (about 10,000 y.b. p.) advances, Riet/ (1912) and Brown (19.5.')) re- August 1985 Crayfishes and Shrimps of Illinois 417 corded localities for O. propinquus and O. indianensis in the Little Wabash River system that almost certainly were based on misidentified O. slannardi (Page 1985). Data collected during the present survey suggest that O. pro- pinquus, O. indianensis, and O. stan- nardi occur allopatrically and that neither O. propinquus nor O. indianen- sis occurs in the Little Wabash River system. Unfortunately, the collections made by Rietz and Brown cannot be located and re-examined. Although all collections of O. stan- nardi that have been made since 1972 are from the Little Wabash River proper, some of the collections of Rietz (1912) and Brown (1955) were from tributaries. Collections of Rietz (1912) were made in Big Muddy Creek between Richland and Clay counties and in Skillet Fork at Wayne City, Wayne County; the former was identified by Rietz as O. propinquus and the latter as O. in- dianensis. Collections cited by Brown (all as O. propinquus but which must have been of O. slannardi) were made in Dismal Creek, 2\ mi N lola. Clay County; Salt Creek, 1 mi SE Effingham, Effingham County; Blue Point Creek, 2'2 mi S Shumway, Effingham County; Lost Fork, I'j mi E Omega, Marion County; branch of Little Wabash River, 1'4 mi SW Trowbridge, Shelby County; and Little Wabash River, 4 mi NE Shumway, Effingham County. Most individuals were found in shal- low riffles composed of large flat stones (Fig. 150). A few were found in deeper water but always in association with stones or accumulations of sticks and other debris. The preference of O. slannardi for rocky riffles is typical of the Propinquus group of Orconecles, the members of which typically hide under stones and among debris. O. slannardi is locally common in the Little Wabash River. Life History.—All seven collections of O. slannardi presently available (from November 1973, Nf)vember 1983, May 1984, and August 1984) contain form I males, and one (May 1984) con- tains two ovigcrous females. All 20 males collected in November, 5 of 1 1 collected in August, and 1 of 25 collected in May are form I. The two ovigerous females collected in May 1984 were 20.7 mm and 28.9 mm CL and carried 124 and 184 eggs, respectively. Eggs average 1.9 mm in diameter. The largest speci- men is a 38.3-mm CL female collected on 22 August 1984; the largest male, from the same collection, is a 30.5-mm form L Orconectes virilis (Hagen) (Fig. 151) Cambarus virilis Hagen 1870 Carnbarus debilis Bundy 1876 Cambarus wisconsinensis Bundy 1876 Cambarus couesi Streets 1877 Description.—Rostrum excavated; moderately long acumen; no median carina; margins thick, straight, barely converging anteriorly, with terminal spines. Carapace flattened dorsoven- trally, with cervical spines; suborbital margin bluntly angular. Areola narrow, constricted anteriorly, narrowest part about 6-7 percent of length. Chelae large, heavily punctate; palm with two rows of tubercles on dorsomesial mar- gin. Form I gonopod terminates in two long, caudodistally directed elements: a sclerotized central projection and a shorter unsclerotized mesial process. Dorsally Hght brown to dark green- brown; fingers of chelae without bright red tips and subdistal black bands. O. virilis occasionally occurs as a blue color morph in Illinois and else- where. Sugar Creek, a tributary of the Sangamon River in McLean County, contains a large number of blue O. virilis. The upper reaches of Sugar Creek are highly modified and receive a large amount of pollution from Bloomingion-Normal. and ihe large number of blue crayfish may somehow be related to the degradation of the stream. Anderson (1975) descrilxxl a 4 1 -mm C;i, female O. virilis from the Kisliwaukie Ri\er at De Kalb. Illinois, that was "chalky white" overall except 418 Illinois Natural History Survey Bulletin Vol.33. ArL 4 i Fig. 151.—Orconecfes virilis. A, dorsal view of carapace; B, dorsal, view of right chela; C, annulus ventralis; D, mesial view of gonopod of form I male; E. lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. August 1985 Crayfishes and Shrimps of Illinois 419 Fig. 152.—Total distribution of Orconectes virilis. The northern limits of the range are un- known. for daikl> pigmented eyes. The closest relatives of O. virilis are O. causeyi, which may be conspecific with O. virilis (Hobbs 1974b), and O. nais. O. fiais is \ery similar to O. virilis morphologically but is distinct serolog- ically (Pryor & Leone 1952). An area of intergradation between O. inrilis and O. nais may be present in southwestern Iowa (Phillips 1980). Distribution and Habitat.—O. I'nilis ranges from New England across south- ern Canada to the Continental Divide in Montana, southeastern Wyoming, and northeastern Colorado, and south to the Ohio River in Ohio, Indiana, and Illinois, the Missouri River in Missouri, and the Platte River in Ne- braska (Fig. 152). The northern limits of its range are unknown (Crocker & Barr 1968), but it occurs farther north than does any other North American crayfish. O. virilis is the most common cray- fish in Illinois and, in some streams, exists in huge populations. It occurs naturally throughout the state except in the .Saline River system and in streams draining the Shawnee Hills and the eastern Coastal Plain (Fig. 153). It is common to abundant over much of its Illinois range, but less so in the lower-gradient streams characteristic of much of western and southeastern Illinois. In the past, its abundance led to its large-scale consumption by humans (e.g., Creaser 1932), but this no longer appears to be true, at least in Illinois. Although absent from the Shawnee Hills, O. virilis occupies Clear Creek and Mill Creek, both of which drain the Ozark Plateaus Province in Union and ' Alexander counties. Habitats in the "Illinois Ozarks" appear similar to those in the Shawnee Hills, but several animals occupy the Ozark Uplift and not the Shawnee Hills, e.g., the snow scorpionfly, Boreus brumalis; the Fig. 153.— Distribution of Orconectes virilis in Illinois: black dots = 1972-1982 collections. 420 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 Fig. 154.—Orconecfes virilis lives in a variety of habitats but usually is found in rocky areas or accumulations of logs and debris such as this logjam on the Little Wabash River, 2.5 mi W Watson, Effingham County, Illinois (8 October 1984). slender madtom, Noturus exilis; the fantail darter, Etheostoma flabellare; and the green water snake, Nerodia cyclopion (Webb, Penny, & Marlin 1975; Smith 1961, 1979). A recent record for O. viriUs in an impoundment in the Saline River sys- tem (Berkel Lake, Saline County, 9 October 1977) is almost certainly the result of a recent introduction. The record is based on two individuals, one of which is small (16.3 mm CL), sug- gesting that the species is reproducing in the lake. In Illinois, O. virilis occupies rocky streams (Fig. 154) and lake margins. It usually occurs in moderate current but is less common in swift riffles (where, throughout much of Illinois, it is replaced by O. propinquus) and in sluggish pools (where O. inunimis predominates). The largest populations are found in areas where large flat stones provide a preponderance of hiding places. In lakes, O. virilis may occur as deep as 9 m (Crocker &: Barr 1968). Cralley (1932) found numerous bur- rows belonging to O. virilis opening just above the water line along the banks of the Kaskaskia Ri\er in Champaign County. Brown (1955) found a few burrows of O. virilis in Illinois and described them as being simple, with an enlarged pocket at the lower end and only one surface opening. Phillips ( 1980) found a burrow in Iowa about 30 cm deep with two terminal chambers and a chimney about 5 cm tall. Hazlett et al. (1974) found O. I'irilis burrowing extensively along a stream and pond in Michigan; burrows frequently had two entrances, often one above the water level. Rarely during the present survey was O. I'irilis found in burrows, and it appears to be much more common for O. virilis to hide under rocks, even in winter, than to burrow; however, as discussed below, o\ igerous females may habiiualh burrow. Life History.—The wide-ranging and often abundant O. i'irilis has been the subject of numerous studies. In both August 1985 Crayfishes and Shrimps of Illinois 421 Table 17.—Frequency of occurrence of form I males, females carrying eggs (ovigerous) and females carrying young, in Illinois collections of Orconectes virilis. 422 Illinois Natural History Slrvev Bulletin Vol. 33, An. 4 in several Ontario lakes (Momot 1978). Momot & Gowlng (1977) found much lower egg counts among high-density populations of O. virilis in Michigan lakes. The number of attached eggs on 402 females (22-35 mm CL) aver- aged only 91.9. These females were smaller than those found in Illinois. Also, fecundity in these lakes was thought to be related to density- de- pendent processes, i.e., as the density increased the number of eggs carried by females decreased (Momot Sc Gowing 1977). Elsewhere, form I males have been found in April and June-October in New England (Crocker 1979), May and August in New York (Crocker 1957), June-August in Ontario (Crocker & Barr 1968), and July-April in Iowa (Caldwell & Bovbjerg 1969). Form I males may be as small as 25 mm CL (Berrill 1978; Weagle 8c Ozburn 1972). Females carrying eggs have been found in April and May in Iowa (Cald- well Sc Bovbjerg 1969; Phillips 1980), April and May in Michigan (Pearse 1910; Momot & Gowing 1977) and New Hampshire (Crocker (1979), May in Rhode Island (Crocker 1979), and May- July in Ontario (Crocker Sc Barr 1968). Females with young attached have been collected in May in Iowa (Phillips 1980) and in June in Maine (Crocker 1979). Some Illinois specimens appear to be among the largest recorded, although Creaser (1932) noted that in Wisconsin the species "frequently attains a size of over eight inches" (about 100 mm CL). The largest Illinois specimens are from reservoirs: a 59.5-mm CL fe- male from Dawson Lake, McLean County, collected on 25 September 1972, and a 59.2-mm CL form I male from Baldwin Lake, Randolph County collected on 2 November 1972. O. mrilis is an agressive crayfish and, with its large size giving it an advantage, ac- tively excludes the often syniopic O. iniDnniis from desirable hiding places (Bovbjerg 1970). Intiaspecific domi- nance was examined by Bovbjerg (1953). The sex ratio among specimens col- lected during the present sur\ey was 1,243 males: 1,1 82 females. Genus Fallicambarus Hobbs FaUicambarus Hobbs 1969 Gonopod with two large terminal elements curved at about 90° angle to shaft; central projection large and bladelike. Ischium of third pereiopod of male with hook. Opposable margin of dactyl of chela with deep conca\ ity. Only F. fodiens of the 14 recognized species of Fallicambarus (Hobbs 1981) occurs in Illinois. F. fodieiis is assigned to the subgenus Creaserinus and is most closely related to F. hedgepethi (see comment below) and F. uhleri (Hobbs 1973). Fallicambarus fodiens (Cottle) (Fig. 155) Astacus fodiens Cottle 1863 Cambarus argil licola Faxon 1884 Description.—Rostrum broad, ex- cavated, with short acumen, lacking marginal spines and tubercles, lacking median carina. Carapace compressed, lacking cervical spines; suborbital margin smooth (not angular). ,\reola obliterated. Chela large, punctate; palm with one to three rows of tubercles on mesial margin; dactyl with deep concavity on basal half of opposable margin. Form I gonopod terminates in two elements, both cur\ ed at an angle of about 90° to the principal axis of the gonopod: a distally rounded sclero- tized central projection and a short proximally thick mesial process with a pegiike tip. Dorsal cx:)loi variable, usu- ally olive brown with dark brown blotches, but often a uniform light brown to rust red. Distribution and Habitat. — F. fo- difns is the most wide-ranging species in its genus, extending, in the Great Lakes (except Lake Superior) and Mis- sissippi Ri\er drainages, from souiiiern Ontario to wesieni Kentuckx (Fig. 156). South of Kentucky it is replaced by the clo.sely related, and probablv conspe- cific (Burr S: Hobbs 1984; H. H. Hobbs August 1985 Crayfishes and Shrimps of Illinois 423 Fig. ^55.—Fallicambarus fodiens. A, dorsal view of carapace; B. dorsal view of right chela, C, annulus ventralis; D, mesial view of gonopod of form I male; E, lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. 424 Illinois Natural History Survey Bulletin \'ol.33.Art. 4 Fig. 156.—Total distribution of Fallicambarus lodiens. The southern limit of its range is un- known. personal communication), F. Iiedge- pethi, which ranges to the Gulf Coast and into Gulf drainages from Georgia to Texas. In Illinois it is known from the Ohio (including the Wabash), Big Muddy, Sangamon, Illinois, and Lake Michigan drainages (Fig. 157). It appears to be absent from western and extreme north- ern Illinois and is rare in central Il- linois. Although difficult to collect because of its burrowing habits, present records suggest that it remains common in southern Illinois in the Ohio, Big Muddy, Saline, and Little Wabash systems. During the present survey. F. fodieiis was collected in central Illinois only on the Sangamon River floodplain at Allerton Park in Piatt County; however, historical records suggest that the species was formerly more common in central and northern Illinois. The northernmost records shown in Fig. 157 are based on a collection at the Il- linois Natural Historv Survey labelled Fig. 157.— Distribution of Fallicambarus fodiens in Illinois. Large circles = 1890-1901 collections; small circles = collections made in the 1950's; black dots = 1972-1982 collections. -Ottawa, LaSalle Co., 111.. Sept. 1901," on Bovbjerg's (1952) study of F. fodiens and O. propinquus in the Lake Michi- gan drainage of southern Cook County, and Bovbjerg's (1952) mention of the presence of F. fodiens in Dead River at Zion, Lake County, also a tributary of Lake Michigan. Brown (1955) found F. fodiens in a woodland pond (Sangamon River system) 2.5 km SW Langley\ille, Christian County. In Illinois, F. fodiens lives on wooded floodplains (Fig. 158) or wheie wooded floodplains formerly occurred. It lives in deep burrows and is in surface waters (usually floodplain ponds) only dining floods: 27 of .'51 collections of F. fodiens made in Illinois since 1972 were made from February through May, the period of late winter-spring flooding, although specimens also ha\ e been caught above August 1985 Crayfishes and Shrimps of Illinois 425 Fig. 158.—Lower Simmons Creek, 5 mi N Golconda, Pope County, Illinois (26 October 1983). Fallicambarus fodiens is common in burrows along the stream banks and, during floods, in the stream. ground in June, July, August, and September. As floodwaters recede and floodplain ponds dry, F. fodiens bur- rows underground. In southern Illinois, chimney-topped burrows on wooded floodplains usually are those of F. fodiens. Stands of river birch (Betula nigra) are especially good indicators of the likely presence of F. fodiens. In Ontario, F. fodiens inhabits marshes, drainage ditches, and ponds (Crocker & Barr 1968). > Form I) Fig. 159.—Size-frequency distribution of Fallicambarus fodiens collected in April in southern Illinois. Table 18.—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Fallicambarus fodiens. 426 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 I Burrows constructed by F. jodiens typically have one to three entrances, often capped by chimneys, leading to an oval resting cavity (Faxon 1885) and leading down from the cavity 30-60 cm is a narrow escape tunnel (Crocker & Barr 1968). Soils having a high clay content are probably necessary for the deep burrows constructed by F. fodiens (Crocker & Barr 1968). Life History.—In a study in southern Cook County, Illinois, Bovbjerg (1952) found juveniles and gravid females in woodland ponds as early as March. By June the adults had disappeared, pre- sumably into burrows, and only young remained in the ponds. By 2 weeks prior to the drying of the pond, all crayfish had disappeared from the pond, either by burrowing or emigrating. Individ- uals were found (one at a depth of 1.2 m) throughout the summer by digging in the dry pond bed down to the water table. The refilling of the pond in the fall was followed by a reinvasion of F. fodiens, which was active in the pond in November under a 5-cm ice cover. F. fodiens can survive out of water (at 20.5° C and 66 percent relative hu- midity) for about 12 hours (Bovbjerg 1952), and overland migrations of F. fodiens on warm (>5°C) humid nights in March-May were docu- mented by Cummins (1921). F. fo- diens is omnivorous, and in northern Illinois specimens, stomach contents included plant fragments (especially grass seeds), insect fragments, crayfish appendages, an isopod, and a sala- mander larva (Bovbjerg 1952). Sexually mature (form I) males occur in March and April in Michigan (Cum- mins 1921) and in April (Table 18), August, and October (Brown 1955) in Illinois. The two form I males col- lected during the present survey had carapace lengths of 42 and 43 mm. In Michigan, females have been found with eggs in March and April and with young in April and May (Pearse 1910; Cummins 1921: Creaser 1931), in Indiana with eggs and young in April (Hay 1896: Williamson 1907). in Ontario with eggs in March and April and young in May (Crocker & Barr 1968), and in Illinois with eggs in Feb- ruary and March and young in Feb- ruary, March, and April (Table 18). Brown (1955) dug a female with at- tached young from a burrow in south- ern Illinois in October. In the present survey, three females with eggs and 24 females with attached young were collected. The females with eggs were 22, 28, and 35 mm CL, with 48, 68, and 162 eggs, respectively. Eggs on the two smaller females, one col- lected each in February and March, averaged 2.2 mm in diameter; those on the largest female, collected in March, averaged 2.5 mm. Seven females col- lected in February and March (30-44 mm CL) had 96-201 (mean, 132.1) young attached. The size-frequency distribution of southern Illinois, April-collected F. fodiens suggests that the species reaches sexual maturity and dies in its second year. Only males over 40 mm CL (second year) among the indi\iduals collected were form I, and all females over 30 mm CL (second year) were carrying young. All smaller males and females were immature (Fig. 159). The sex ratio among the combined April samples was 44 males: 42 females. Genus Cambarus Erichson Carnbarus Erichson 1846 Bartonius Ortmann 1905 Gonopod with two large terminal elements curved at about 90° angle to shaft; central projection large and bladelike. Ischium of third pereiopodof male with hook. Opposable margin of dactyl of chela without deep concavity. Cambarus contains about 77 species (Hobbs 1981), of which four occur in Illinois. Two of the Illinois species (lenebrosus and rusticiformis) belong to the subgenus Erebicambarus, one (dio^enes) to Lacunicambarus. and one (robuslus) to Punclicambarus. The wide-ranging C. diogenes appears to be a complex of species (Hobbs 1974b), August 1985 Crayfishes and Shrimps of Illinois 427 Fig. 160— Cambarus tenebrosus. A, dorsal view of carapace; C, annulus ventralis: D, mesial view of gonopod of form I male; form I male; F, lateral view of gonopod of form II male B, dorsal view of right chela; E, lateral view of gonopod of 428 Illinois Natural History Survey Bulletin Vol. 33, An. 4 more than one of which may occur in Illinois, as discussed in the species account below. Cambarus tenebrosus Hay (Fig. 160) Cambarus bartoni tenebrosus Hay 1902 Cambarus bartoni laevis Faxon 1914 Description.—Rostrum broad, shal- lowly exca\ated, lacking median carina; margins slightly converging anteriorly, lacking terminal tubercles or spines; acumen short. Carapace dorsoventrally flattened, cervical spines absent (north- ern population) or present (southern population); suborbital margin gently angular. Areola moderately wide, nar- rowest part about 13-14 percent of length, with at least four punctations across. Chela large, heavily punctate; palm with one or two rows of tubercles on mesial surface. Form I gonopod terminates in two elements, both curved at an angle of more than 90° to the principal axis of the gonopod; a distally rounded sclerotized central projection, and a large tapering to broadly rounded mesial process. Dorsally rust red over- all, with abdomen slightly browner than carapace. In a discussion of the distribution and phylogeny of species of Cambarus, Hobbs (1969) depicted the range of C. (Erebicambarus) laevis as occurring mostly north of the Ohio River and that of the consubgeneric C. tenebrosus as being south of the Ohio River. Hobbs (1974b) separated the two species on the relative width of the areola ("at least 4 punctations across narrowest part" in C. tenebrosus; "no more than 3 punctations across narrowest part" in C. laevis) and the shape of the mesial process of the gonopod ("broadly rounded distally" in C. tenebrosus; "strongly tapered, often subacute" in C laex'is). Populations of the subgenus Erebi- cambarus occurring in southern Il- linois have been referred lo as C. bartoni brevis (Rietz 1912), C. bartoni laei'is (Brown 1955), C. laevis (Hobbs 1968, Fig. 161.—Total distribution of Cambarus tenebrosus. Fig. 162, brosus in I lections. —Distribution of Cambarus tene- linois; black dots = 1972-1982 col- August 1985 Crayfishes and Shrimps of Illinois 429 1969, 1972a; Page 1974b: Peck & Lewis 1977), C. ornalus (Peck & Lewis 1977), and C. teuebrosus (Peck & Lewis 1977). It now appears that the correct name for the southern Illinois populations is C. teuebrosus. Populations extend across the Shawnee Hills of southern Illinois, into the Cumberland River drainage of western Kentucky, and into the general range of C. teuebrosus with no notice- able mor|)hological discontinuities. In \'ermilion County in east-central ^^ "K^uX^^jti Fig. 163.—Spring emptying into Big Creek in Hardin County, Illinois (8 October 1984). This and other springs in the Shawnee Hills provide the best habitat in Illinois for Cambarus tene- brosus. 430 Illinois Natural History Survey Bulletin Vol. 33, An. 4 Fig. 164 —Cambarus rusticiformis. A, dorsal view of carapace; B, dorsal view of right cfiela; C, annulus ventralis; D, mesial view of gonopod of form I male; E. lateral view of gonopod of form I male; F, lateral view of gonopod of form II male. August 1985 Crayfishes and Shrimps of Illinois 431 Illinois is a population of Erebicam- barus discovered in 1977. It differs from C. tenebrosus in southern Illinois by having a narrower areola and by lacking cervical spines (although tubercles often are present). No form I males have been found. If C. laevis is a valid species, the X'ermilion County population is morphologically and geographically (jide Hobbs 1969) referable to that taxon. However, the absence of dis- continuities in the geographic variation in morphology among populations assignable to the C. tenebrosus-C. laevis complex suggests a lack of reproductive isolation among populations and that only one species is recognizable. In- dividuals in the northern part of the range of the species have narrower areolae, lack cervical spines, and ap- parently have more tapered mesial processes on the gonopods. H. H. Hobbs, Jr. (personal communication 1983) suggests that "when adequately studied, laevis will prove to be a syno- nym of tenebrosus." Pending such a study, all Illinois populations are re- ferred to as C. tenebrosus. Distribution and Habitat.— C. tene- brosus occurs in central and south- eastern Indiana, the Vermilion River system in east-central Illinois, the Shawnee Hills of Illinois, western and central Kentucky, central Tennessee, and northern Alabama (Fig. 161). It appears to be absent in southwestern Indiana. In Illinois and elsewhere, C. tene- brosus occupies rocky springs and spring-fed headwaters and creeks. It hides under stones, sometimes forming shallow excavations in the stream bed. Brown (1955) cited "small piles of mud at the edge of rocks" as an indicator of the presence of the species. Its affinity for spring-fed headwaters often results in its occupancy of caves. In southern Illinois, C. tenebrosus is restricted to the Ohio River drainage of Hardin, Pope, Johnson, and Union counties (Fig. 162). It is most common in Big Creek, a moderate-sized stream in Hardin County fed by numerous Fig. 165.—Total distribution of Cambarus rusticiformis. Fig 166— Distribution of Cambarus rustici- formis in Illinois; black dot = 1975 collection. 432 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 springs (Fig. 163). Elsewhere in Illinois it has been found only in Willow Creek (Vermilion River system) in Vermilion County, Illinois (Fig. 162), where it presumably is distributionally contig- uous with the Indiana population. The Illinois and total ranges of C. lenebrosus are quite similar to those of the zigzag salamander (Plethodon dorsalis) although the salamander has dispersed into Clear Creek, a tributary of the Mississippi River, in Alexander, Union, and Jackson counties (Smith 1961). C. tenebrosus, first recorded in Illi- nois (as C. bartoni brevis) by Rietz (1912), probably has essentially the same distribution now as it had prior to the arrival of man. However, the water table of southern Illinois has been lowered within historic times, and some springs may have disappeared or at least are smaller than before, reducing the habi- tat available for C. tenebrosus. Such reductions in habitat have caused some spring-inhabiting fishes (Phoxinus erythrogaster, Rhinichlhys atratulus) to disappear from southern Illinois (Smith 1979). Life History.—Prins (1968) studied C. lenebrosus in a stream system in Ken- tucky and found it to be most common in constant-temperature headwaters. Most adults were found in burrows or under large rocks, but young and sub- adults were found in open-stream environments. The diet consisted pri- marily of vascular plants, especially fallen leaves from the riparian habitat, and filamentous algae. Other food items included detritus, aquatic insects, and crustaceans. Hatching occurs in July and August after eggs have been carried by the female for at least 25-30 days. Growth continues throughout the year, including winter, and sexual maturity is reached at about 20-22 months and 10 mm CL. Mating occurs in winter and spring. Mature Irmales oviposit in June and July at about 22-23 months and probably again at 34-35 months. Egg counts on seven females (44-52 mm (;L) ranged from 186 to 266. Maximum CL was found to be about 65 mm and maximum longevity about 36-38 months. The sex ratio among 3,235 specimens was 1 male: 1.03 females. No form I males or females carrying eggs or young were present among the 40 collections of Illinois C. tenebrosus made during the present survey (all months except September). Brown (1955) reported collecting form I males in August and October and a female with attached young in October. All of Brown's collections were made in Hardin and Pope counties, Illinois. A 46.6-inm CL female from Big Creek is the largest Illinois specimen. Cambarus rusticiformis Rhoades (Fig. 164) Cambarus rusticiformis Rhoades 1944 Description.— Rostrum narrows anteriorly; margins elevated, concave, and tipped with large knobs; acumen moderate, ele\ated anteriorly; median carina lacking. Carapace dorso\entrally flattened, with large cervical spine, small suborbital angle. Areola mod- erately wide, narrowest part 16-18 per- cent of length. Chela large. hea\ily punctate; palm with single row of flattened tubercles along mesial surface. Form I gonopod terminates in two ele- ments, both directed caudalK at an angle greater than 90° to the principal axis of the gonopod: a distally rounded, sclerotized central projection and a large distally tapering mesial process. Distribution and Habitat. — C. rusti- ciformis occupies the Cumberland River drainage and the Barren Ri\er system (Green Ri\er drainage) of Ken- lucky and Tennessee (Bouchard 1976) and, judging from one Illinois collec- tion, the lower Ohio River (Fig. 165). Two juvenile males (11.7 and 14.6 mm CL) were collected in a rockv backwater of the Ohio Ri\er near Lock and Dam 53, 4 km NE Olmsted, Pulaski County, Illinois, (Fig. 166) on 16 July 1975. Several later attempts to find C. rusti- ( iformis at the same and other sites on the lower Ohio Ri\er were imsuccessful. August 1985 Crayfishes and Shrimps of Illinois 433 r ^nL^V^^^f^!"^^/;"^ diogenes. A, dorsal view of carapace: B, dorsal view of right chela- Pnr^T^^LT 'f ",°' "^^n" ^'"^ °* aonopod of form I male: E, lateral view of gonopod oiform I male: F. lateral view of gonopod of form II male 434 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 Fig. 168.—Total distribution of Cambarus diogenes. The Ohio River site is about 67 river km (42 miles) below the mouth of the Cumberland River. The two specimens may represent one of the westernmost natural populations of the species, they may have been waifs captured during a downstream dispersal, or they may have been transplanted as the result of human activities. C. rusticiformis has been introduced into the Paint Rock River (Tennessee River drainage), Alabama (Bouchard 1976). Collected in the Ohio River with C. rusticiformis were 12 Orconectes placidus, a species with a similar overall distribution. Elsewhere, C. rusticiformis lives in rocky runs and adjacent pools and riffles of fast-flowing creeks and small rivers. Other ecological characteristics are un- known. Cambarus diogenes Girard (Fig. 167) Astacus fossor Rafinesqiie 1817. (Name suppressed by Iiiieniaiional Commission on Zooligical Nomenclature, Opinion 522, 1958.) Cambarus diogenes Girard 1852 Cambarus nebrascensis Girard 1852 Cambarus obesus Hagen 1870 Cambarus Diogenes var. Ludoviciana Faxon 1885 Description.—Rostrum broad, ex- cavated, with short acumen, lacking marginal spines or tubercles, lacking median carina. Carapace compressed, lacking cervical spines; suborbital margin angular. Areola obliterated. Chela large, heavily punctate; palm with two or three rows of tubercles on mesial surface; dactyl with broad shal- low concavity on basal half of oppos- able margin. Form I gonopod termi- nates in two elements, both directed caudally at an angle slightly more than 90° to the principal axis of the gonopod: a distally rounded, sclerotized central projection and a slender mesial process. Dorsal color varies from uniformlv Fig. 169.— Distribution of Cambarus diog- enes in Illinois; blacl< dots = 1972-1982 col- lections. August 1985 Crayfishes and Shrimps of Illinois 435 ;-..^:^i*s^: -^^^2^"^'.::^^^:-:^^ ^^P^**lj.^^ Fig. 170.—Chimney-topped burrows of Cambarus diogenes on the floodplain of a slough, 2 mi N Pleasant Grove. Johnson County, Illinois (9 July 1972). red brown (with thorax lighter red) to green with red outHning abdominal segments, tips of chelae, and margins of rostrum. Marlow (1960) examined geographic variation in C. diogenes and concluded that only two subspecies could be rec- ognized: C. d. ludovicianus in the Lake Ponchartrain drainage and southern Louisiana and C. d. diogenes through- out the rest of the range of the species. However, according to Hobbs (1969, 1974b), C. diogenes is a complex of forms, and C. d. ludovicianus probably is more widespread than Marlow thought. Illinois specimens show tre- mendous variation in the shape of the carapace and rostrum and in coloration. If several taxa presently are included in "C. diogenes." more than one may be found in Illinois. The only other de- scribed species in the subgenus Lacuni- cambarus are C. acanthura, restricted to the Tennessee and Alabama drain- ages of Tennessee, Alabama, and (ieoi- gia (Hobbs 1981) and C miltus from Baldwin Clounty, Alabama (Filzpatritk 1978b). Dislribulion and Habitat. — C. diog- enes ranges along the Atlantic Pied- mont and Coastal Plain from New Jersey and Pennsylvania south to cen- tral Georgia, in Gulf drainages from Georgia to eastern Texas, and north in the Mississippi and Great Lakes basins to western New York and Penn- sylvania, Michigan, Wisconsin, Min- nesota, and North Dakota. The western- most populations occur in the Platte Ri\er drainage of Colorado and Wy- oming (Fig. 168). C. diogenes occurs statewide in Il- linois (Fig. 169) and is generally com- mon. It is abundant in the Shawnee Hills and on the Coastal Plain of south- ern Illinois, less common in northern and western Illinois. Its burrowing habits preclude easy collecting, and consequently, the number of (olle( lions (Fig. 169) does not adetjuateh reflect its abundance in Illinois. C. diogenes first was recorded from Illinois by Hagen (1870), who described the spec imens (ollected at Lawn Ritlge (Marshall County), Belleville, and Fvanston as a new species, Cambarus obesus. Forbes' (1876) only comments on "('.. obesus" were: "Very tommon. The largest in the state." The burrowing l)ehavioi of C. diog- 436 Illinois Natural History Survey Bulletin Vol. 33, An. 4 Table 19—Frequency of occurrence of form I males, females carrying eggs (ovigerous), and females carrying young, in Illinois collections of Cambarus diogenes. August 1985 Crayfishes and Shrimps of Illinois 437 Fig 171.-Cambarus robustus. A. dorsal view of carapace; B, dorsal view of right chela- C, annulus ventralis; D, mesial view of gonopod of form I male; E. lateral view of gonopod oftorm I male; F, lateral view of gonopod of form II male. 438 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 were collected from February through July (except in May), and in October and November (Table 19); Brown (1955) found form I males in Illinois from March through September, except in July. The primary peak of occurrence is in February and March, and a second- ary peak occurs in November (Table 19). Elsewhere, matings have been observed in March, April, May (Hargitt 1890; Hay 1896; Evermann Sc Clark 1920), and October (Williams & Leonard 1952). In Louisiana, form I males were found in March, April, and October (Penn &: Marlow 1959) and in Georgia in Jan- uary, April, and October (Hobbs 1981). Ortmann (1906) recorded form I males in Pennsylvania from March through November. Among the C. diogenes examined by Marlow (1960), form I males were present in every month ex- cept January, but were most prevalent in April and September. Although the occurrence of form I males suggests that mating in C. diogenes may occur in almost any month, activity probably peaks in late winter-early spring (Feb- ruary-April) and in fall (September- November). Ovigerous females have been found in January (Bundy 1877), April (Hay 1896; Evermann &: Clark 1920: Creaser 1932), and May in Indiana (Williamson 1907); in April and May in Pennsyl- vania (Ortmann 1906) and Iowa (Phil- lips 1980); in April (Williams & Leonard 1952) and May in Kansas (Harris 1900); from late March to May in Wisconsin (Graenicher 1913); and in September in Georgia (Hobbs 1981 ). Marlow ( 1960) examined o\igerous females from March (1 female), April (19), May (17), and October (I). Females carry young in May and June in Pennsylvania (Ortmann 1906) and in Indiana (Wil- liamson 1907; Evermann &: Clark 1920), and in June in Michigan (Creaser 1932). In Illinois, females were found to carry eggs from January through Ma\, except Fig. 172.—Total distribution of Cambarus robustus. Fig. 173,— Distribution of Cambarus robus- tus in Illinois. The two records (Adams and Macon counties) are from Faxon (1885) and Rietz (1912). August 1985 Crayfishes and Shrimps of Illinois 439 in February, and young in May and June (Table 19). Because of the difficulty in obtaining adult C. diogenes, few females with eggs or young attached were collected in the present survey. Five females (43.7-56.4 mm CL) collected in April and May were carrying 183-518 (mean, 295.2) eggs. The highest number of eggs was on the smallest female. Eggs varied in diameter from 2.1 to 3.0 mm, and unlike other crayfishes examined dur- ing the present survey, in C. diogenes egg diameter has a significant corre- lation (r = 0.97) with length of the fe- male: egg diameter = -1.280 + 0.077 CL. A 48.5-mm CL female collected in, Johnson County on 19 May 1973 had 606 early-instar young attached; a 47.2- mm CL female collected in Christian County on 8 June 1979 had 45 late-instar young attached. Two females collected in Indiana in 1906 were carrying 87 and 215 young (Williamson 1907). Cambarus robustus Girard (Fig. 171) Cambarus robustus Girard 1852 Description.—Rostrum moderately broad, shallowly excavated, lacking median carina; margins converge an- teriorly, lack terminal tubercles or spines; acumen moderately large. Cara- pace dorsoventrally flattened, with large cervical spines; suborbital margin angular. Areola moderately wide, nar- rowest part about 12-17 percent of length. Chela large, heavily punctate; palm with two rows of tubercles on mesial surface; base of fixed dactyl with deep dorsal and ventral imjjres- sions. Form I gonopod terminates in two elements: a distally rounded, sclero- tized central projection curved at an angle of more than 90° to the principal axis of the gonopod, and a large gently tapering mesial process curved at about 90° to principal axis. Dorsally green brown. Distribution and Habitat.—C. ro- bustus ranges from Connecticut and southern Ontario south to northwestern North Carolina and eastern Kentucky and west to Michigan and central In- diana (Fig. 172). It occupies rocky areas of fast-flowing streams, and sometimes is found in rocky lakes and ponds (Crocker &: Barr 1968; Berrill 1978). The only Illinois records (Fig. 173) are from the late nineteenth and early twentieth centuries. Faxon (1885) re- ported C. robustus from Decatur (Macon County, Sangamon River sys- tem), and Rietz (1912) reported four specimens from Quincy (Adams County, Mississippi River drainage). Although the Quincy specimens cannot be located, the Decatur specimens are in the U.S. National Museum (USNM 63291) and unquestionably document the former presence of the species in the state. C. robustus is rare even as far west as Indiana (Eberly 1955), and the Illinois collections must have repre- sented relict populations left over from a more widespread distribution in a cooler postglacial climate. Many streams today around Quincy are spring-fed, and considerable effort was expended in unsuccessful attempts to locate surviving populations of C. robustus in what appear to be suitable habitats. The Sangamon River, flowing through Decatur, was badly polluted in the early part of this century (Jewell 1920), is now impounded by Lake De- catur, and appears unlikely to support a population of C. robustus. Life History.—Crocker (1957) re- ported form I males in New York col- lections made from April through Octo- ber and observed copulation in October and May (the latter in a lamprey trap). Form I males have been collected in Pennsylvania in May and July-No- vember; the smallest was about 63 mm total length (about 32 mm CL) (Ort- mann 1906). In southern Ontario, form I males were 25-55 mm CL (Berrill 1978). Ovigerous females have been found in July in New York (Crocker 1957) and Pennsylvania (Ortmann 1906); the Pennsylvania female, 84 mm total length (about 42 mm CL), carried 228 eggs. Females with young attached 440 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 have been observed in April and August in New York (Crocker 1957). Young feed on immature insects; adults feed mainly on aquatic plants (Greaser 1934). CONSERVATION Unfortunately, it appears that Macro- brachium ohione and Cambarus ro- biistus already have disappeared from Illinois. C. robustus may have dis- appeared from Illinois naturally rather than because of man-induced changes in the environment. Our earliest records indicate that it was relict in Illinois, presumably having been reduced from a more widespread distribution to small areas near Quincy and Decatur. Else- where it inhabits cool to cold streams and apparently was reduced in warm postglacial Illinois (probably during the Xerothermic period) to small popu- lations that subsequently were imable to perpetuate themselves. A7. ohione probably disappeared because of the extensive modification and degradation of Illinois' large rivers. Especially harmful loM. ohione would have been the loss of aquatic vegetation associated with increased siltation, channeliza- tion, and chemical pollution. Unless protective measures are taken, several more species of native decapods are likely to disappear from Illinois. The following four species have re- stricted ranges in the state and are threatened with extirpation. They should receive as much protection as possible by our adding them to the Illinois List of Endangered and Threat- ened Species and protecting their habitats. The distribution of Orconectes in- dianensis has been reduced dramatically in Illinois as a result of severe pollution and other major modifications of the Saline River system. Remaining popu- lations are restricted to the South Fork of the Saline River in Gallalin, John- son, Pope, Saline, and Williamson (ounties, Robinette (Eagle) Creek in (.allaiin Comity, and Honey Creek in ilavdin C^ounty. Elsewhere the species occurs only in a small area of Indiana. Orconectes kentitckiensis occurs only in Big, Hosick. and Peters creeks in Hardin Comity, and although the dis- tribution of this species in Illinois probably has changed little historically, its persistence is threatened by modifica- tions of the riparian habitats in Hardin County. Trees have been remo\ed from many miles of stream bank, resulting in less shading of the stream and higher water temperatures. Water has been diverted in ever-increasing amounts from its natural drainage for use in agriculture. The lowered water table is reducing the size of Big Creek. Else- where, O. keutuckiensis occurs only in a small area of northwestern Kentucky. Orconectes placidus is common throughout much of Kentucky and Tennessee, but in Illinois it maintains a sizeable population only in Big Creek in Hardin County and is threatened by the same stream modifications threat- ening O. kenluckiensis. Orconectes lancijer is restricted in Illinois to Horseshoe Lake in .\lexander County. Although under the supervi- sion of the Illinois Department of Con- ser\ation, the lake has been severely modified in recent decades, and much of it now is surrounded by agricultural fields ami grasses. If the unusual biota of Horseshoe Lake is to persist, subsequent modifications of the lake shoidd be allowed only after carefid consideration of their iin ironmental inipaci. Else- where O. lancifrr occurs imcoinmonly ihoughout the former Mississippi Em- bayment. ACKNOWLEDGMENTS I am extremely indebted to Mr. Craig W. Ronto for the execution of most of the figures appearing in this pidilica- tioii and lo Dr. H. H. Hobbs, jr.. for laxononiic comisel, distributional in- formation, and a thorough review of the manuscript. I am indebted also to B. M. Burr and R. W. Bouchard for taxonomic (inmsel during the study, to J. .\. Beatty, P. W. August 1985 Crayfishes and Shrimps of Illinois 441 Smith, G. C. Sanderson, and C. Burnett for helpful reviews of the manuscript, to Bernice P. Sweeney for typing the manuscript, and to the following col- leagues and friends for assistance with field work and/or providing specimens: J. Allen, R. V. Anderson, N. A. Ashford, S. S. Birky, L. Boscarine, J. K. Bouse- man, J. A. Boyd, R. A. Brandon, A. R. Brigham, W. U. Brigham, B. Britton, J. Brower, K. Brummett, J. Buerger, B. M. Burr, P. A. Burr, M. K. Butcher, R. W. Clegern, L. E. Cordes, A. Covich, F. Cruse, K. S. Cummings, D. M. Cur- rent, R. Cygan, L. R. Davis, D. Dufford, W. H. Ettinger, R. A. Evers, J. Fairchild, J. Grady, R. Grosser, L. R. Hanebuvg, W. Herridon, R. Hite, R. W. Horner, M. Hoverstein, D. L. Johnson, T. M. Keevin, W. L. Keller, M. King, M. Klutho, L. Kolczak, J. Langbein, E. A. Lisowski, E. L. List, K. Lubinski, C. Maier, J. C. Marlin, W. Mathis, W. Matsunaga, R. L. Mayden, J. McNur- ney, R. L. Moran, M. A. Morris, V. L. Morris, D. A. Nolan, L. D. O'Bryant, S. D. Ogle, D. Osterbur, E. M. Page, M. L. Page, V. M. Page, P. J. Paladino, P. Parrillo, G. Pefrates, I. Polls, A. Pulley, F. Putz, M. Ratke, M. E. Retzer, L. R. Richart, R. Rogers, C. W. Ronto, K. Russell, D. Sallee, M. W. Sanderson, R. T. Schaaf, R. Schacht, D. W. Schem- ske, P. W. Smith, S. Smith, L. J. Stan- nard, R. Stinauer, J. Sublett, L. B. Sulo- way, C. S. Swanson, B. Taubert, R. Thom, R. Thomas, J. Thomerson, J. Tranquilli, W. J. Tucker, R. Twillman, J. D. Unzicker, W. S. Vinikour, W. Ward, T. Wayne, D. W. Webb, J. Webb, M. J. Wetzel, F. Winter, L. A. Woodrum, R. D. Wrisberg, and L. S. Zimmer. LITERATURE CITED Aiken, D. E. 1969. Ovarian maiuration and egg laying in the crayfish Orconecles iiirilis: influ- ence of temperature and pholopcriod. Canadian Journal of Zoology 47:931-935. Anderson, R. V. 1975. An albinistic crayfish, Orconectes virilis (Hagen, 1870) (Decapoda, Astacidea). Crustaceana 29:314. Berrill. M. 1978. Distribution and ecology of crayfish in the Kauartha region of southern Ontario. Canadian Journal of Zoology 56:166- 177. Black, J. B. 1966. Cyclic male reproductive activ- ities in the dwarf crawfishes Cambarellus shu- feldti (Faxon) and Cambarellus puer Hobbs. American Microscopical .Society Transactions 85:214-232. . 1972. Life history irotes on the crawfish Orconecles lancifer. Louisiana Academy of Sciences Proceedings 35: 7-9. Bouchard, R. W. 1976. Geography and ecology of crayfishes of the Cumberland Plateau and Cumberland Mountains, Kentucky, Virginia, Tennessee, Georgia, and Alabama. Part 1. The genera Procambarus and Orconecles. Part II. The genera Falticambarus and Cambarus. Pages 563-605 in J. W. Avault. Jr., ed.. Fresh- water crayfish. Louisiana Slate Division of Continuing Education. Baton Rouge. . 1977. Threatened and recently extinct crayfishes of the western United States (Deca- poda: Astacidae and Cambaridae). Association of Southeastern Biologists Bulletin 24(2): 38-39. and H. W. Robison. 1980. An inventory of the decapod crustaceans (crayfishes and shrimps) of Arkansas with a discussion of their habitats. Arkansas Academy of Science Proceed- ings 34:22-30. BovBjERG. R. \'. 1952. Comparative ecology and physiology of the crayfish Orconecles propin- quus and Cambarus fodiens. Physiological Zoology 25:34-56. . 1953. Dominance order in the crayfish Orconecles virilis (Hagen). Physiological Zoology 26: 173-178. 1970. Ecological isolation and competi- tive exclusion in two crayfish (Orconecles viri- lis and Orconecles tmmunis). Ecology 51:225- 236. Boyd, J. A., and L. M. Page. 1978. The life his- tory of the crayfish Orconectes kenluckiensis in Big Creek, Illinois. American Midland Naturalist 99:398-414. Braun, E. L. 1950. Deciduous forests of eastern North America. Blakiston Co., Philadelphia. Broad, A. C and J. H. Hubschman. 1963. The larval development of Palaemonetes kadiaken- sis M. J. Rathbun in the laboratory. American Microscopical Society Transactions 82: 1 85- 1 97. Brown, P. L. 1955. The biology of the < rayfishes of (eniral and southeastern Illinois. Doi toral Dissertation. I'niveristy of Illinois. I'rbana- C;hainpaign. 168 p. . 1956. A new crayfish of the genus Orco- necles from Illinois (Decapods. Astacidae). American Midland Naturalist 56:163-167. 442 Illinois Natural History Survey Bulletin Vol.33. ArL 4 . 1959. Coinpaiisoii of habitats of twotiay- fi.shcs of the genus I'rocambarus in Illinois and Louisiana. Illinois Academy of .Scienie Trans- arlions ,52:104-108. Bryan, C. F., F. M. Truesdale. and D. S. Sabins. 1975. Annual lepoil, a limnological survey of the Alchafalaya Basin. Louisiana Cooperative Fisheries Research I'nil, .School of Forestry and Wildlife Management, Louisiana State University, Baton Rouge. BuNDY, VV. F. 1877. On the Cambari of northern Indiana. Vcadeiny of Natural Sciences of Phila- delphia Pioceedmgs 29:171-174. . 1882. A list of the Crustacea of Wisconsin with notes on some new or little known species. Wisconsin Academy of Science, Arts, and Let- ters Transactions 5:177-184. Burr, B. M.,andH. H. HoBBS,Jr. 1984. Additions to the crayfish fauna of Kentucky, with new locality records for Cambareltus shujeldiii. Kentucky Academy of Science Transactions 45:14-18. Busch, K. H. D. 1940. Embryology of the cray- fish, Cambaru.s rusticus Cirard. Doctoral Dis- sertation. The Ohio State University, Colum- bus. Caldwell, M. J., and R. \'. Bovbjerg. 1969. Nat- ural history of the two crayfish of northwestern Iowa, Orconecles x'irilis and O. immunii. Iowa Academy of Science Proceedings 76:463-472. Capelli, G. M. 1982. Displacement of northern Wisconsin crayfish by Orconecles rusticu.s (Girard). Limnology and Oceanography 27: 741-745. , and J. F. Capelll 1980. Hybridization between crayfish of the genus Orconecles: mor- phological evidenie (Decapoda, Cambaridae). Crustaceana 39:121-132. , and J. J. Magnlison. 1983. Morphoe- daphic and biogeographic analysis of crayfish distribution in northern Wisconsin. Journal of Crustacean Biology 3: 548-564. , and B. L. iVUinjal. 1982. Aggressive inter- actions and resource coiripetition in lelation to species displacement among crayfish of the genus Orconecles. Journal of Crustacean Bi- ology 2: 186-492. Chambers. C. L, J. F. Payne, and M. L. Kennedy. 1979. Geographic variaton in the dwarf cray- fish, Cambarellus puer Hobbs (Decapoda, CaiTibaridac). Crustaceana 36: 39-55. Clark, T. H.. and C, W. .Stearn. 1960. Ihe ge- ological evolution of North America. Ron.ild Press Co.. New York. Crallev. J. W. 1932. .\ sur\ey of the crayfishes of Champaign Comny, Illinois. Master's I'hesis. University of Illinois, Urbana-ChampaigiL CRKA.SER, E. P. 1931. The Michigan decapod crustaceans. Michigan .\cadem\ of Science, Arts and Letters Papers 13:257-276. . 1932. rhe decapod Crustacea of Wiscon- sin. Wisconsin .\cadeiriy cjf Science. .-Vrts and Letters Tiansactions 27:321-338. 1933. .\ freshwater shrimp for the tropical aquarium. Aquarium 1:261-262 1934. A biological survey of the Raquette watershed. \'. The larger Crustacea of the Rac)uette watershed. Supplement to the 23rd Annual Report. New York Conservation De- partment, Biological Survey 8:158-163. . and \. I. Ortenblrcer. 1933. The deca- pod crustaceans of Oklahoma. I'niversity of Oklahoma Biological Survev Publication 5(2):13-47. Crocker. D. W. 1957. The crayfishes of New York State (Decapoda. Aslacidae). New York State Museum and Science Seivice Bulletin 355. . 1979. The crayfishes of New England. Biological Society of Washington Proceedings 92: 225-252. . and D. W. Barr. 1968. Handbook of the crayfishes of Ontario. I'niversity of Toronto Press. Cummins. H. 1921. Spring migration in the cray- fish. Cambarus argilhcola Faxon, .\merican Microscopical .Si5ciely Transactions 40:28-30. Darnell, R. M. 1958. Food habits of fishes and larger invertebrates of Lake Pontchartrain. Louisiana, an estuarine conmiunity. Institute of Marine Sciences Publications. I'niversity of Texas 5:353-356. Eberly, W. R. 1955. Summary of the distribution of Indiana crayfishes, including new state and county records. Indiana .\cadem\ of Science Proceedings 64:281-283. Engle. E. T. 1926. Crayfishes of the genus Cam- barus in Nebraska and eastern Colorado. Bu- reau of Fisheries Bulletin 42:87-104. EvERMANN. B. W.. and H. W. Clark. 1920. Lake Maxinkuckee, a physical and biological survey. Indiana Departnient of C>inser\ation Publica- tion 7. EvERS, R. A., and L. M. P.«;e. 1977. .Some ususual natural areas in Illinois. Illinois Natural His- tory Survey Biological Notes 100. Faxon, W. 1885. .\ revision of ihe .\slacidae(Part 1. The genera Catnbarus and Aslncus). Museum of Comp.iiative Zoology Memoirs. Harvard College H)(4):l-186. . 1890. Notes on North .\merican cray- fishes, family ,\stacidae. I'.-S. National Museum Proceedings 12:619-634. . 1914. Notes on the cra\ fishes in the I'niied States National Museum .nul the Museum of Comparative /.oology, with description of new species and subspecies to which is apptMided a catalogue of the known species and subspe- cies. Museum of Comp.iiative Zoologv Mem- oirs, Harvard C:ollege 40(8): 351-427. August 1985 Crayfishes and Shrimps of Illinois 443 Fenneman. N. M. 1938. Phvsiograpln of enslern I'nilcii Stales. McGraw-Hill Book Co., New York. FiDi.AR, M. M. 1948. Phybiogiaphy o( the lower Wabash \'alley. Indiana Depailnieni of Con- servation. Division of Geology, Bulletin 2. FiSK. H. M. 1944. Geological investigation of the alluvial valley of the lower Mississippi River. Mississippi Ri\er Commission Report. FiTZPATRiCK. J. F., Jr. 1967. Ihe Propinquus group of the craw'fish genus Orconecles (Deca- poda: .-Xstacidae). Ohio Journal of .Sdcnce 67:129-172. . 1968. A new crawfish of the genus Orco- necles from Iowa. American Midland Naturalist 79:507-512. 1978a. A new crawfish of the subgenus Girardiella, genus Procnmbarii.s from north- west Arkansas (Decapoda. Cambaridae. Bi- ological Society of Washington Proceedings 91:533-538. 1978b. \ new burrowing irawlish cjf the genus Cnmbanis from southwest Alabama (Decapoda, Cambaridae). Biological Society III Washington Proceedings 91:748-755. . 1983. A revision of the dwarf crawfishes (Cambaridae, Cambarellinae). Journal of Crustacean Biology 3: 266-277. , and B. A. Laning. 1976. A new dwarf crawfish (Decapoda: Cambaridae: Cambarel- linae) from southwest Alabama and adjacent Mississippi. Biological Society ol Washington Proceedings 89:137-146. Fleming. L. E. 1969. I'se of male external genita- li( details as laxonomic characters in some species of Palaemonetes (Decapoda, Palae- monidae). Biological Society of Washington Proceedings 82:443-452. F'lint, R. F. 1971. Glacial and Quaternary ge- ology. John Wiley & Sons, Inc., New York. Forbes. S. .\. 1876. List of Illinois Crustaiea, with descriptions of new species. Illinois Museum of Natural History Bulletin 1 ( 1 ): 3-25. , and R. E. Richardson. 1908. The fishes of Illinois. Illinois State Laboratory of Naiuial History. 357 p. + atlas of 103 m;ips. Fowler, H. W. 1912. The Caustacea o( New Jer- sey. New Jersey Stale .Museum .Viinual Rcpiiri for 1911:29-6,50. Francois, D. D. 1959. The crayfishes of New Jer- sey. Ohio Journal of Science 59:108-127. Graenicher. S. 1913. .Some noles on the habits and distribution of Wisconsin crawlishes. Wisconsin Natural History Society Bulleiin 10:118-123. Grow, L. 1981. Buirowing beha\joi ui ihcday- fish, Cambarus diogene.s diogenes Girard. Animal Behavior 29:351-3.56. GuNTER, C;. 1937. Observations on the river shrimp, Macri>bracliium ohionis (Smith). American Midland Naturalist 18:1038-1042. Hagen H. a. 1870. Monograph of the North American Astacidae. Illustrated Catalogue of the Museum of Comparative Zoologv, Harvard Colle.ge 3. HargittC:. W. 1890. Some habits of the ciay-fish. .Vmerican Monthly Microscopical Journal 11:111-114. Harris, J. ,\. 1900. .\nnotated catalogue of the crayfishes of Kansas. Kansas University Quai- lerly, .Series A, 9:263-274. . 1902. Distribution of Kansas crayfishes. Kansas I'niversity Science Bulletin 1:1-11. Hav, O. p. 1882. Notes on some fresh-water Crus- tacea, together with descriptions of two new species. .American Naturalist 16:143-146. 241- 243. Hay. W. p. 1896. The crawfishes of the state of Indiana. Pa.ges 476-506 in 20th Annual Report of the Department of Geology and Xiitural Resources of Indiana. Hazlett. B., D. Rittschof, and D. Ri benstein. 1974. Behavioral biology of the ciaylish Or- conecte.s virilis. I. Home Range, .\merican Midland Naturalist 92:301-319. ' Hedgpeth, J. W. 1947. Freshwater shrimp. Texas Game and Fish 5:14-15. . 1949. The North .American species of Macrobrachium (river shrimp). Tex.is Journal of Science 1(3): 28-38. Herrick, F. H. 1896. The American lobster: a study of its habits and development. U.S. Fish Commission Bulletin 15:1-252. HoBBS, H. H., Jr. 1942. The crayfishes of Florida. University of Florida Publication, Biological Science .Series 3(2). . 1962. Notes on the affinities of the mem- bers of the Blandnigu section of the crayfish genus Procambarus (Decapoda, .\stacidac). Tulane Studies in Zoolog:>' 9:273-293. . 1968. Crustacea: Malacostraca. Pages K-l-K-36 in F. K. Parrish. Keys to water cjuality indicative organisms (southeastern I'nited Stales). Federal Water Pollution Cionirol \i\- ministi.iiion. U.S. DeparlmenI of the Interior. 191)9. On the disiiibuiion and [ihylogeny of the ciayfish genus Cambarus, Pages 93-178 in P. C. Holt, R. L. Hoffman, and C. W. II.nl, Jr., eels. Ihe disli ibulional history of the bioia of the Southern .\ppalac hi:ins. Part I: Invciie- brates. Virginia PoKlechinc Insniule Rc-se.uch Division Monogiaph 1, . 1972a. Crayfishes! Aslacidae)of norih ,uid middle .\merica. U.S. Environmental Piolec- lion ,\gencv Biota of Freshwater Ecosystems Identification Manual 9. 444 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 . 1972b. The subgenera of (he crayfish ge- nus Procambarus (Deapoda: Astacidae). Smith- sonian Contributions to Zoology 117. 1973. New species and relationships of the members of the genus Fallicambarus. Biolog- ical Society of Washington Proceedings 86: 461 - 482. 1974a. Synopsis of the families and genera of crayfishes (Crustacea: Decapoda). Smith- sonian Contributions to Zoology 164. 1974b. A checklist of the North and Mid- dle American crayfishes (Decapoda: Astacidae and Cambaridae). Smithsonian Contributions to Zoology 166. 1977. The crayfish Bouchardina robisom. a new genus and species (Decapoda, Cambari- dae) from southern Arkansas. Biological Society of Washington Proceedings 89: 733-742. . 1979. A new crayfish from the Ouachita River basin in Arkansas (Decapoda: Cambari- dae). Biological Society of Washington Pro- ceedings 92: 804-8 1 1 . 1981. The crayfishes of Georgia. Smilh- HuNER. J. V. 1977. Observations on the biology of the river shrimp from a commercial bait fishery near Port Allen. Louisiana. Pages 380- 386 m Thirty-first Annual Conference of the Southeastern Association of Fish and Wildlife Agencies Proceedings. Jewell. M. E. 1920. The quality of water in the Sangamon River. Illinois State Water Survey Bulletin 16:230-246. Langlois, T. H. 1935. Notes on the habits of the crayfish, Cambarus rusticus Girard, in fish ponds in Ohio. American Fisheries Society Transactions 65:189-192. Lewis, J. J. 1982. Systematics of the troglobitic Caecidotea (Crustacea: Isopoda: Asellidae) of the southern interior low plateaus. Brim- leyana 8:65-74. -. and T. E. Bowman. 1981. Thesubterran- sonsian Contributions to Zoology 318. . and P. H. Carlson. 1983. Distocambarus (Decapoda: Cambaridae elevated to generic rank, with an account of D. crocken. new species, from South Carolina. Biological Soci- ety of Washington Proceedings 96: 420-428. -, and A. G. Grubbs. 1982. Description of a new troglobitic crayfish from Mexico and a list of Mexican crayfishes reported since the pub- lication of the Villalobos Monograph (1955) (Decapoda, Cambaridae). Texas Memorial Museum Bulletin 28:45-50. -, and L. J. Marchand. 1943. A contribu- tion toward a knowledge of the crayfishes of the Reelfoot Lake area. Tennessee Academy of Science Journal 18:6-35. HocuTT, C. H., R. F. Denoncourt, and J. R. .Stauffer. Jr. 1978. Fishes of the Greenbrier River, West Virginia, with drainage history of the central Appalachians. Journal of Bioge- ography 5(1): 59-80. HoLTHUis, L. B. 1952. A general revision of the Paiaemonidae (Crustacea Decapoda Natantia) of the Americas. II. The subfamily Patae- moninae. Allan Hancock Foundation Occa- sional Paper 12. HoRBERG, L. 1945. A major buried valley in east central Illinois and its regional relationships. Illinois C;eological Survey Report 106: 349-359. HiiBSCHMAN. J. H., and J. A. Rose. 1969. Palar- monetes kadiakni.sis Rathbun: Post embrvonic growth in the laboratory (Decapoda, Palae- monidae). Crustaceana 16:81-87. ean asellids (Caecidotea) of Illinois (Crustacea Isopoda: Asellidae). Smithsonian Contribu- tions to Zoology 335. Lowe, M. E. 1956. Dominance-subordinance re- lationships in the crawfish Cambarellus shu- jeldtu. Tulane Studies in Zoology 4:139-170. . 1961. The female reproductive cycle of the crayfish Cambarellus shufeldtii: the influence of environmental factors. Tulane Studies in Zoology 8:157-176. Luce, W. M. 1933. A survey of the fishery of the Kaskaskia River. Illinois Natural History Sur- vey Bulletin 20:71-123. Marlow, G. 1960. The subspecies of Cambarus diogeties. .\merican Midland Naturalist 64: 229-250. Matthes, F. E. 1933. The Pleistocene diversion of the Mississippi River across Crowlev's Ridge, southeast Missouri. Science (New Series) 77: 459-460. McCoRMiCK, R. N. 1934. Macrobrachiiim oluoius. the large freshwater shrimp. Indiana .\cadeniy of Science Proceedings 34:218-224. Meehean, O. L. 1936. Notes on the freshwater shrimp Palaemoneles paludosa (Gibbes). American Microscopical Society Transactions 55:433-441. Meredith, W. G., and F. J. Schwartz. 1960. Maryland crayfishes. Maryland Department of Research and Education. Educational Series 46. MoMOT, W. T. 1967. Population dvnamics and productivity of the ciavfish. OKonecles iinhs. in a marl lake. American Midland Natiiralisi 78:55-81. . 1978. .\nnual production and produc- tion biomass ratios of the crayfish. Orconecles I'lrilis. in two norihcrn Ontario lakes, .\merican Fisheries Society Transactions 107:776-784. August 1985 Crayfishes and Shrimps of Illinois 445 , and H. GowiNC. 1977. Production and population dynamics of the crayfish Orconectes virilis in three Michigan lakes. Fisheries Re- search Board of Canada Journal 34: 2041-2055. Newcombe, C. L. 1929. The crayfishes of West Virginia. Ohio Journal of Science 29:267-288. Nielsen, L. A., and J. B. Reynolds. 1975. Fresh- water shrimp natural food for pond fishes. Farm Pond Harvest 9(2): 8,9.24. , and . 1977. Population character- istics of a freshwater shrimp. Palaemoneles kadiakensis Rathbun. Missouri Academy of Science Transactions 10 & 11:44-57. Ortmann, a. E. 1906. The crawfishes of the State of Pennsylvania. Carnegie Museum Memoirs 2: 343-523. . 1931. Crawfishes of the southern Ap- palachians and the Cumberland Plateau. Car- negie Museum Annals 20:61-160. Pace, L. M. 1974a. The life history of the spottail darter, Elheostorna squamueps, in Big Creek, Illinois, and Ferguson Creek, Kentucky. Illinois Natural History Survey Biological Notes 89. . 1974b. Aquatic Malacostraca recorded for Illinois, with notes on their distributions and habitats within the state. Illinois State Academy of Science Transactions 67: 89-104. . 1975. The life history of the stripetail darter, Elheostorna ketinicottt, in Big Creek, Illinois. Illinois Natural History Survey Bio- logical Notes 93. 1985. A new crayfish of the genus Orconectes from the Little Wabash River sys- tem of Illinois (Decapoda: Cambaridae). Bi- ological Society of Washington Proceedings 98:564-570. , and B. M. Burr. 1973. Distributional records of the crayfishes Cambarellus puer, C. shufeldtii, Procambarus gracilis, P. viae- viridis, Orconectes lancifer, O. bisectus, and O. rusticus. Kentucky Academy of Science Trans- actions 34:51-52. Pearse, a. S. 1910. The crawfishes of Michigan. Michigan State Biological Survey Publication 1:9-22. Peck, S. B., and J. J. Lewis. 1977. Zoogeography and evolution of the subterranean invertebrate faunas of Illinois and southeastern Missouri. Natural Speleological Society Bulletin 40:39- 63. Penn, G. H. 1942. Observations on the biology of the dwarf crayfish, Cambarellus shufeldtii (Faxon). American Midland Naturalist 28: 644- 647. . 1943. A study of the life history of the Louisiana red-crawfish, C.atnbarus clarkii Girard. Ecology 24:1-18. -. 1950. The genus Cambarellus in Louisi- . 1952. The genus Orconectes in Louisiana (Decapoda, Astacidae). American Midland Naturalist 47: 743-748. . 1956. The genus Procambarus in Louisi- ana (Decapoda, Astacidae). American Midland Naturalist 56: 406-422. . 1959. An illustrated kev to the crawfishes of Louisiana with a summary of their distribu- tion within the state (Decapoda, Astacidae). Tulane Studies in Zoology 7: 3-20. and J. F. Fitzpatrick, Jr. 1962. Inter- specific competition between crawfishes. Amer- ican Zoologist 2: 436. , and . 1963. Interspecific competi- tion between two syinpatric species of dwarf crawfishes. Ecology 44: 793-797. -, and H. H. Hobbs, Jr. 1958. A contribu- tion toward a knowledge of the crawfishes of Texas (Decapoda, Astacidae). Texas Journal of Science 10:452-483. , and G. Marlow. 1959. The genus Cam- barus in Louisiana. American Midland Nat- uralist 61:191-203. Phillips, G. S. 1980. The decapod crustaceans of Iowa. Iowa Academy of Science Proceedings 87:81-95. Prins, R. 1968. Coinparative ecology of the cray- fishes Orconectes rusticus and Cambarus tene- brosus in Doe Run, Meade County, Kentucky. International Revue Gesamten Hydrobiologie 53:667-714. Pryor, C. W., and C. A. Leone. 1952. Serological comparisons of astacuran Crustacea. Biological Bulletin 103:433-445. Reimer, R. D. 1969. A report on the crawfishes (Decapoda, Astacida) of Oklahoma. Oklahoma Academy of Science Proceedings 48: 49-65. , K. Strawn, and a. Dixon. 1974. Notes on the river shrimp, Macrobrachium ohione (Smith) 1874, in the Galveston Bay System of Texas. American Fisheries Society Transac- tions 103:120-126. Rhoades, R. 1944a. The crayfishes of Kentucky, with notes on variation, distribution, and de- scriptions ol new species and subspecies. Ameri- can Midland Naturalist 31:111-149. . 1941b. Furthei studies on distribution ana (Decapoda, Astacidae). American Midland Naturalist 44:421-426. and taxonomy of Ohio crayfishes, and the de- scription of a new subspecies. Ohio Journal of Science 44:95-99. RiECEL, J. A. 19.59. The systematics and distri- bution of crayfishes in California. Calilornia Fish and Game 45:29-50. RiET/., N. M. 1912. Ecological relations of the crawfishes of Illinois. B. S. Thesis. Lniversity ol llliiKiis, I'rbana-Champaign. 446 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 .S.viiTH. D. G. 1981. Evidence for hybridization between two crayfish species (Decapoda: Cam- baridae: Orconectes) with a comment on the phenomenon in Cainbarid crayfish. American Midland Naturalist 105:405-407. Smith, P. W. 1957. An analysis of post-Wisconsin biogeogiaphy of the Prairie Peninsula region based on distributional phenomena among terrestrial vertebrate populations. Ecology 38(2): 205-218. . 1961. The amphibians and reptiles of Illi- nois. Illinois Natuial History Survey Bulletin 28:1-298. . 1971. Illinois stieams: a classification based on their fishes and an analysis of factors responsible for disappeaiance of native species. Illinois Natural History Survey Biological Notes 76. . 1979. The fishes of Illinois. University of Illinois Press, I'rbana. Strenth, N. E. 1976. A review of the systemaiics and zoogeography of the freshwater species of Palaeinonele.s Heller of North America (Crus- tacea: Decapoda). Smithsonian Contributions to Zoology 228. Tack, P. I. 1941. The life history of the crayfish Cambarus imriiunis Hagen. American Midland Natuialist 25:420-446. " Teller, J. T. 1973. Preglacial (Teays) and early glacial drainage in the Cincinnati area, Ohio, Kentucky, and Indiana. Geological Society of America Bulletin 84(1 1): 3677-3688. Threinen, C. VV. 1958a. Wisconsin crayfish. Wisconsin Conservation Bulletin 23(7):13-15. . 1958b. A suininary of observations on the commercial harvest of crayfisli in northwestern Wisconsin with notes on the life history of Orconectes viritis. Fish Management Division of the Wisconsin Conservation Deparlmeni Miscellaneous Report 2:1-14. Truesdale, F. M., and W. J. Mermilliod. 1979. The river shrimp Macrobrachium ohionc (Smith) (Decapoda, Palaemonidae): its abund- ance, reproduction, and growth in the Atcha- falaya River basin of Louisiana. I'.S.V. Crusta- ceana 36: 61-73. TtiRNER. C. L. 1926. The ciaylishes of Ohio. Ohio State Universilv Bulletin 30:14,5-195. Unger, P. A. 1978. Natural history- invenion of Colorado. 3. The crayfishes (Crustacea: Cam- baridae) of Clolorado. University of Colorado Museum, Boulder. \'an Deventer. W. C. 1937. Studies on the biology of the crayfish Cambarus propinquus (Girard). Illinois Biological .Monogographs 15(3»:l-67. \'illalobos, a. 1983. Crayfishes of Mexico (Crus- tacea: Decapoda) (Translation by H. H. Hobbs, Jr.) Baba Barkha Nalh Printers. New Delhi. India). VioscA, P., Jr. 1931. The bullhead, Ameiurus melas calulus. as a dominant in small ponds. Copeia 1931:17-19. . 1939. Where to fish in Louisiana for craw- fish. Louisiana Conservation Review 8(1):I7- 18. . 1953. .\ll about crawfish— life history and habits. Louisiana Conservationist 5(6|: 3-5. Wayne, W. J. 1952. Pleistocene evolution of the Ohio and Wabash \alle\s. Journal of Geology 60: 575-585. Weagle, K. \'.. and G. W. Ozbirn. 1972. Obser- vations on aspects of the life historv of the cray- fish. Orconettes virihs (Hageni, in imnhwest- ern Ontario. Canadian Journal of /oology 50: 366-370. Webb, D. W., N. D. Penny, and J. C. .Marlin. 1975. The Mecoplera. or scorpionflies. of Il- linois. Illinois Natural Hisiorv Survev Bulletin 31:250-316. White, F. A. 1949. Preliminary notes on the breed- ing season of Palaemoneles kndiakcnsis Rath- bun in the Baton Rouge .\rea. Louisiana .Acad- emy of Sciences Proceedings 12:71-74. Williams, \. B. 1954. Speciaiion anddisiribution of the crayfishes of the Ozark Plateaus and Ouachita Provinces. l'niversit\ of Kansas Sci- ence Bulletin 36:803-918. , and .\. B. Leonard. 1952. Lhe cia\fishes of Kansas. l'niversit\ of Kansas Science Bulle- tin 34:961-1012. Williamson, E. B. 1907. Notes on the crayfish o{ Wells County, Indiana, with the description of a iww species. Pages 749-763 i>i .\nnual upon. Department of Cieologv and N'atur.i! Resources of Indi.ina. Wii.i.MAN, H. B.. aiul J. C. Frvk. 1970. Pleisuxene sn.itigiapby of Illinois. Illinois Stale Geologi- cal Sur\ev Bulletin 94. INDEX •147 ncanthura. Caxibaru.s, 435 aumlhurus. Miurnbriu liiuiu. 356 aiuliis, C:/unharu\. 376 aculus, Cambaru.s btaniiingii. 376 acutus. Pxocambanis, 376 alnbametisii. Orconcctcs, 390 antrorum. Pnlaemonetes, 359 argillicota. Canibarus, 335, 422 A.slacus. 422 jodtens. 422 fossor. -i'i-i wi.scon.siiii'>isis, 335, 417 larrinu.'i, Ma< robrat htuiii, 356 cau.seyi, Onoiintes, 419 Clarkii, Cambarus. 381 clarkii. Procambarus. 381 Conservation, 440 louesi, Canibarus, 417 Crayfish Life Cycle. 343 Creaserinus, 335 cuevachicae, Procambarus acutus. cummingi, Palaemoneles, 359 376 Barbicambarus, 362 barloni, Catribarus, 428 Barlonius, 426 biseclus. Orconecles, 338 biacki, Cambarellus, 368 blaudingu. Procambarus. 376 blandingii. Procambarus acutus. 376 Boiu hardma. 362 brevis, Camharus bartoni. 335, 428 Cambarellinae, 335 Cambarellus. 335, 363 fc/ac*), 368 diminutus, 368 /cs/ifj, 368 ninae. 368 pufr, 366 schmilti. 368 shufeldtii. 363 /cva/H(i, 368 C:ambaridae, 335, 362 Cambarinae, 335 Canibarus. 335, 426 acanthura, 435 af!((uj, 376 argitlicola, 335. 422 bartoni, 428 ferci'ii. 335, 428 /flpi^w. 428 tenebrosus. 428 blandingti acutus. 376 C/ar/(H.'381 couesi, 417 rfc6i7(i. 417 diogenes. 434 var. Ludoviciana. 434 ludovicianus, 435 jaxonii, 400 gracilis, 370 /H'gH<.v. 335. 376 tenebrosus, 428 viae-i'iridis, 378 turi/i.s, 417 debilis. Canibarus. 417 Derapoda. 335 diminulus, Cambarellus, 368 diogenes, Canibarus, 434 Dirigicambarus, 335, 363 Distocambarus, 362 Erebuurnbarus, 335, 428 exiUpes, Palaemoneles, 335, 360 Extraliniital .Species and Studies, 336 Fallicambarus. 335, 422 fodiens, 422 hedgepethi. 422 ulileri, 422 Faxonelta. 362 faxonii. Canibarus. 400 Faxonius. 386 immunis pedianus. 388 fodiens. .-istai us, 422 fodiens. Fallicambarus. 422 forceps, Orconectes, 412 fossor, Astacus. 434 Freshwater Shrimp Life Cycle, 345 Geological History of Illinois, 339 Girardielta. 335, 371 gracilis, Canibarus, 370 gracilis, Procambarus, 370 Historical Studies of Illinois Decapods, 335 harrisoni. Orconectes, 338. 396 hedgepethi, Fallicambarus, 422 Hohbseus, 362 holthuisi. Palaemoneles, 359 hubbsi, Canibarus, 338 /iy/as, OrfOfiff(«, 338 illinoiensis, Orconectes, 386 immiiMi,?, Canibarus, 388 immunis. Orionectes. 388 immunis \ pedianus. Orconectes, 390 indianensis, Canibarus, 394 indianensis, Orconectes, 394 /ou'ac'i.siA. Orconectes, 406 juvenilis, Canibarus. 412 juvenilis, Orconectes. 412 kadiakensis. Palannonetes, 359 kentuckiensis, Orconectes. 396 Key to Decapods of Illinois. 348 Key to Illinois Oayfishcs Based on Form I M.iles. 353 Key to Illinois Spec ies. 345 l.aiuniuimbarus, 335, 435 448 lancifer, Orconectes, 400 laei'is, Cambarus, 428 lan'is, Cambarus bartoni, 428 lancifer, Cambarus. 400 lesliei, Cambarellus, 368 liberorum. Procambarus. 371 limosus, Orconectes, 412 Ludoviciana, Cambarus diogenes var., 434 ludovicianus, Cambarus diogenes, 435 luteus, Orconectes, 338 Macrobrachium, 335, 356 acanlhurus, 356 carcinus, 356 ohione, 356 oljersii. 356 medius, Orconectes, 338 millus, Cambarus, 435 r!a«, Orconectes, 338. 419 Natural History, 343 r!f6raicfn5«, Cambarus, 434 ninae, Cambarellus, 368 obesus, Cambarus, 335, 434 ohione, Macrobrachium, 356 Ohionis, Palaemon, 356 oljersii, Macrobrachium, 356 Orfonpf/«, 335, 386 alabamensis, 390 bisectus, 338 fOUiiC)!!, 419 forceps, 412 harrisoni, 338, 396 /i5)/a5, 338 illinoiensis, 386 immunis, 388 immunis x pedianus, 390 indianensis. 394 lowaensis, 406 juvenilis, 412 kentuckiensis, 396 lancijer, 400 limosus, 412 /u(fU5, 338 medius, 338 riflii, 338, 419 peruncus, 338 placidus, 404 propinquus, 406 punctimanus, 338 putnami. 412 quadruncus, 338 rhoadesi, 390 riiiiifuj, 412 sloanii, 396 spinosus, 412 stannardi. 415 tricuspis, 338 validus, 390 wiriiii, 417 wrighti, 394 orna(ui, Cambarus, 429 Ortmannicus, 369, 376, 378 Palaemon, 356 Ohionis, 356 sattei, 356 Palaemonetes. 335, 358 an^rori^m, 3.59 (ummin^i. 359 Mi7ipci,'335, 360 holthuisi, 359 kadiakensis, 359 paludosus, 358 /pxanui, 359 Palaemonidae. 335, 356 paludosus, Palaemonetes, 358 Pandicambarus, 335, 368 Paracambarus, 369 Pacijasticus. 362 pedianus, Faxonius immunis, 388 peruncus, Orconectes, 338 placidus, Cambarus. 404 placidus, Orconectes, 404 Procambarus, 335, 369 acij/ui, 376 acutus, 376 cuevachicae, 376 blandmgii, 376 blandingii. 376. 378 c/arAn, 381 gracilis. 370 liberorum, 371 reimeri, 371 troglodytes, 335 inaeviridis, 378 propinquus, Cambarus, 406 propinquus, Orconectes, 406 pufr. Cambarellus, 366 Puncticambarus, 335 punctimanus, Orconectes, 338 putnami, Orconectes, 412 quadruncus, Orconectes, 338 reimeri, Procambarus. 371 rhoadesi. Orconectes. 390 robuslus, Cambarus, 439 ruslicijormis, Cambarus, 432 ruj()fui. Camfcaru5. 412 ru.sii;i, Palaemonetes, 359 tricuspis. Orconectes. 338 Troglocambarus. 362 troglodytes, Procambarus, 335 uhleri, Fallicanibarus, 422 validus, Orconectes. 390 I'lae-viridis, Cambarus, 378 viaei'tridis, Procambarus, 378 viritis, Cambarus, 417 viritis, Orconectes, 417 !i'i.sfo>i.su!f«si.v, Cambarus, 335, 417 wrightt. Orconectes. 394 Zoogeography of Native Illinois decapods. 339 WtT/.KI.. Fauii 1).. B.S., Trihnuai Awtslanl CIrossktt. I.okkiI':. W.S.. Junto j Tn hnutil A\\t\lant Cl'.MMiN<;s, Kkvin S,. M.A., }umT Trt/tttmil AyMstant Irish. Jjn^RFV A., B.S.. Junior Tri Ituunl .\\\i\lnnt Pk\(ITH.i,i. Pamh.a A.. U.S.. Juixuti Tnhnunl .Iwixtant Sandbkrc;. SiitRRi I... B.S,. Junun Trthuual .Iwislant Smkrman. RKNt->. B.S., Junun Trthnual AsMstniil IiMAN. Mahv( ARCH,. B.S.. Juutoi Trtiiiiual .t.wislanl V'ANDrRAii. (ii.KNiJV C... Jututi} Trthuunl Assntant SECTION OF WILDLIFE RESEARCH SANDtRsoN. (,|KN C. Pill).. WthHiff Sf)r( uili^l and Urnil Bki.i.roM':. Frank i... Sc.D.. IVtlilUfr Sfiftialist and Prinapal S( irnlisl CrRABKR, Ri<:hard R.. Ph.D.. Orntl/tolofii.si and Hriiit ipa( St irnlisi hmrTiltt\ Andfrson, Wie.i.iam I... M.S., Wildlifr hcolo^ist Edwards. VVh.iiam R.. Ph.D., IVildliff E(olo^i.\l Grabfr. Jf an \V.. Ph.D., (>rnilholoiii.\i Han.son. Harold C, Ph.D., Wiidlijr Sf>rtiaii\i Nixon, Chari ks M.. M.S., iVildlifr t.toloRist OocHKAN. \V, \V.. Jr., B.S,, Awtxmtr Wildliff .Spntalist Havf-Ra. Stepmkn p., Pri.D,, Awtuuilr IVitdlifr h(ologi\t Larkin. RtiNAi.n P., Ph.D.. Awottalr Witdhlf hi olo^ixt QiINF,. Doi'fii.As B,. Ph.D., Avsoiialr Riof)h\Mi i.\l Warner. Richard E-. Pll.D.. Avuxinlf WiUHifr tUohtfinl \Vf_stfmkier, Ronald I,.. M.S., .-{wotiatr W'lldhfr Emiogist VV Jkan D. .Schoknecht, Ph.D.. Ass*,- iiale l*ri>frs.sor of l.ife Siirnirs. Indiana State ('nwersily. Terrr Haute. Faunisiit Surveys and ln.sccl Idenlificaiion, Roderick R. Irwin. B.A., Chita^o. Illinois: Wildlife Research. Nanc^ I. Bi'RLEY, Ph.D., Assoiiate Professor of EioUtffy. Etholotry. and Etoluliou. ( 'niversity of Illinois: RoBjRT L. Jonfis, Ph.D.. I'rofrssor of Soil A/oirrn/oiry and Eiolu^y. I'nnersily of IHinois: Willjid D, KlimNli.i. Pii D.. /*r.;- frssor of /.oology and Dirntor of (:oof>erative Wildtifr Researth. Southern Illinois I'nnersily: .hkI Norman D. Levink, Ph.D.. Pro- fessor of I'rtrrinary f*arasiloloicv. I'elerinary Rrsran h. and Zoology, .iiid Dim lor of the Crnler for Human EioUi^s. I'nivrrsily of Illi- nois: Enlomology, Nelda Alge:r, Ph.D., Assmiate professor of (•enrlus and Prvrlopmrnl. t 'nn-rrsily of lllinms: Robert I.. NIftcai.f. Ph.D.. I'rofrssor of Biology and Hr.seardi Professor of Enloinoloizy . i'mversity of Illinois: and Gn bfrt P Waidbai fB, Pii O , /'f'>- frMor of Entomology. I'nivrr.Mly of Illinois. (48.816—3M—8-85) Some Publications of the ILLINOIS NATURAL HISTORY SURVEY BULLETIN Volume 33, Article 1.— Dipiera, or True Flies, of Illinois. I. Tabanidae. L. L. Pechuman, Donald W. Webb, and H. J. Teskey. April 1983. 122 p. \'<)lume 33, Article 2.—An Annotated Bibliog- raphy of the Illinois Herpelological Literature 1960-1980, and An Updated Ciheiklist of .Species of the State. Michael A. Morris, Richard .S. Funk, and Philip W. .Smith, April 1983. 15 p. Volume 33, Article 3.— 12.') Years of Biological Research 1958-1983: A Symposium. George Godfrey, Chairman; John Bouseman, William Edwards. Kenneth Robertson, and Robert Zewadski, eds., Proceedings of the 125th An- niversary Symposium of the Natural History Survey. 1985. 200 p. BIOLOGICAL NOTES 120.— Recreational Fishing in the Kankakee River. Illinois, Robert J. Graham, R. Weldon Larimore, and William F. Dimond. June 1984. 13 p. 121.— Bibliography of Illinois Vegetation. Paul C;. Risser. August 198-1. 51 p. 122.—The Life History of the Mud Darter, Elheosloma asjjrigptie, in Lake Greek, Illinois. Kevin S. Gummings, James M. Grady, and Brooks M. Burr. Decemliei 1984. 16 p. 123.—The Life History of the Shorthead Red- horse. Moxosloma miurolrpidolitin. in the Kankakee River Drainage. Illinois. .Michael J. Sulc and Thomas M. Skelly. 1985. 16 p. 124.— Illinois Birds: Vireos. Jean W. Graber. Richard R. Graber. and Ethelyn Kirk. 1985. 52 p. SPECIAL PUBLICATION Number 2.—Landsca|)e Ecology: Directions and Approaches. Paul G. Risser. James R. Karr. and Richard T. F. F'orman. November 1984. 18 p. CIRCULAR 49.—The Dunesland Heritage of Illinois. Her- bert H. Ross. August 1963 (Reprinted .Mav 1974). 28 p. 51.— Illinois Trees: Selection, Planting, and Citre. J. Cedric Carter. March 1977 (Third printing). 123 p. 52.— F'ertiliiing and Watering Tiees. Dan Neely and E. B. Himelick. Decemln-r 1971 (Third printing). 20 p. 54.—Corn Rootvvoiin Management in Ciinning Sweet Corn. W. H. Luckmann. J. T. Shaw, D. E. Kuhlman. R. Randell. and G. D. LeSar. March 1975. 10 p. 55.—Observing. Photographing, and Collecting Plants. Kenneth R. Robertson. August 1980. 62 p. MANUSCRIPTS High (|ualily manuscripts dealing with any aspect of natural history will be considered for publi- cation ill one of the Illinois Natural History .Survey publication series: Bulletin, Binlogiral S'olrs. Circular, and Spr< ml fublicalion. I'he author need not be an employee of the Survey, but if not. will be required to pay priming costs. Manuscripts must follow the reccmimendations of the third edition of the Council of Biology t'.ditors Style Manual except that journal namc-s in the Liteiature Citi-d section are to be s|)elled out in full. Fhe Survey expects to publish only one or two manuscripts by non-Survey authors yearly. Two or more outside referees reconnnend each manuscript submitted for publiralion in ihc^BuI- letin series before it is accepted. Please send three copies of manuscripts to Ix- considered for publication to Office of the Chief, Illinois Natural History Survey, 607 East Peabody Drive, ("hampaign, Illinois 61820. /,/.v( oj available publications mailed on request No charge is made for publications of the Illinois Natliral History Survey. A single copy of most ptd)lications will be sent free to anyone requesting it until the supply becomes low. Cx)sily pub- lications, more than one copy of a pid)licati