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- \oulltrTJi I srulinii tfir i*re<,i
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 Levinf, ti.i. Ph.D.. AssiMiatr hnlomologisl Stefffv. Kevin. Ph.D., .-tsso
Rfinboi D, Kfti'Rah a,. PhD . Awnnate Aqualu Binl<>si\t Mil I FR. t-RFUERic D.. Ph.D.. .issistant Entomoiuf^ist. f.xtrnston
Ross. Phji iPCF., Pii.D . Aystxtatf Aqunlu /iioi(>iii\l Wei, I.kstlr, Pii.D . Assistant Prnfrwumal Srimttst
Bavi.fv. Pftkr B., Ph.D.. Ay-^nlanl Aqualu liuilof'isl Wein/ilri , Richard A,, Pw.li., .•isststant hntomolof^tst, f.xteniirtn Hfnfbrv. Mtchaki. S., Ph.D.. Awistanl Aqunlu Builoi^i\l /avaifta, l.iis, Ph.D.. Assistant Frofrsnonal \timtttt Hicks. Randai.i,. Ph.D.. As'.islani Aquniu Hinloiiist Heim, Charles. C;.. M.S., AssiMiatr Supportn-r Sfirnlist Horns, Wh.i.iam H.. Ph.D.. AsM\ianl Aqualu Bioloiiist koGAN. Jenny, M.S., Associalr Supportn-r St irntist I.l'BiNSKl, Kennfiii S., Ph.D.. Ay\islant Aqualu Binlo^isl WiusoN. Jean Ci.. B.A., .4sswiatr Supportn-r Stirnttst Starnf.s, Wavnf. PhD-. A\si\laul Aqualu Biolotiist .\i bif. JoAnn, Assistant Suppitrtur S
W'arrfn. (.arv 1. , B.S.. AsM.Uanl Supptntivf SnrtUnl l.EpAR. CfERAl D. Assistant Supfmrtn-r Surnlist Ai'sTFN. Doi'f.l.AN, M.S.. }u>u(n I'Tiilrwinnal SiifnliU McC.iFFFN. MiiTON K,. Jr.. ,\I.S., .4ssistant Supportn-r Stimtisl Hoof,, M u:h a F-I. M.S.. Juuun l>Ti>frss,„nal St irnli.yl HaYER. Karen. .M.S.. Assistant Suppifrtwr Si irntist HiftFRT. Bfveri.f.y. Junior l*ii)fe\Mi)iial Surnlist Roberts. Stephen J.. B.S., .issislant Supportn-r Scirntist KwAK. Thomas |.. M.S., Junmr Pmfrwinunl \iifttlt\t .Swoeford. David. Ph.D., Assistant Suppojtn-r Siirnlisl Pfrrv, l.ANCF-. M.S.. Juniur I'rnfr.ynitnal Si it'Ulisl Wai.sh, i.Al'RA. M-S., .-tssisiant Suppinln'r S* imtisl SoBASKl. .StfPHES v.. B.S., luni'ir I'rulrwional Siirntist Da/ey. Doyi e. Junior Profrssumat Stirnlisl .Sons. tii<;FNK. Junior PrnfrsMotml S< irnlist Post, SrsAN. Junior Suppurln'r Stirntisl Iazik. Pamf-1-A p.. M.S.. Junuir P^ofewional Siifnti\l (Carpenter. Sherla I.., \..\., Tnhnical Assistant Dovvi,iN(.. David, M.S., Tri imnal Assistant i'.jssY. I.At'RiE, B.S., Trihnual .issistant
twiN(., Kathar^n I... B.S,, Tnhniinl Awistanl PeteR-s. Brenda M.. Trihnual Assistant KoPHFiMAN, Jfffrf\. M.S.. Tfi liniinl Awistanl Sherman, Renee. B.S., Trihnual A ssisiani I.voN, Dai.f. M.S.. Tnhnunl Assislnnt WiisMANN, Janet I... B.S.. Trihnual Assistant Nf.wman. Barky t.. B.S., Tr-ihnual Assislnnl WiTiiROW. Jane. Trt hnual Assistant A.s.sisiant .ScHMiTTi.FH. Crak. D . M.S., Trifiuuat Assi\lant Morris. Marilyn. M.S.. yumor Trihnual Wai.nfr. Ri'iH. Tr< hnnal A\si\lnnt Mc(;iiRE. .Michael. M.S.. Junior Trrhnual Axststani
{^I.Ai'S.SF.N. Ji'I.iF,. B.S., Junior Trihnual .-tssnlnnt CtRlDER. DntolliCii. Tran.si-ribinfi Srtrrtary, l:\lrnMtttl C^RAWFORD, LoI'Ann, B.S,. Junior Tri linual A.wnlanl Lrdman, Diane. (.Irrk-Typist III
(;ros.sf.tt. Lorrik, MS., Junior Tfihnuat .-ixMslitnt 1 RAiB, Joan. (Irrk-T\pist III DfQi'fnnf, Barbara, yuM/or Tnhnifal Assistant S/KonxiNsKl. .\\.\c.f,(.lrrk-Typisl II Das, Daviu, E.S.. Junior Tn hniial A\\i.\lnnt BaI'CHMAn. I'homas \.. (iraduatr Hrsranh As.sislanl K1FID.S, Robert, H.S.. Juuioj Tnlinnnl A<
Mavfr. (jiRl.STiNF, B.S-, Junior Trt hnual AsM\tanl KiELDiNc, Dennis J., (iraduatr Trathin^ Assistant 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 SECTION OF FAUNISTIC SURVEYS AND INSECT WiKF. I.VNN, M.S., Junior Tri hnii at .Assistant IDENTIFICATION
LaBergf. Wali A( e t.. Ph D,. Insri-t Taxonomisl and Hrad
SECTION OF BOTANY AND PLANT PATHOLOGY Brigham. Warren l\, Ph.D . Insrft Taxonomist
Pa(.f. Iamrencf M.. Ph D . Fish Taxonomisl (iRUNWAi.u, Ci.Ai'N. I'h.D., Botanist ami Hratt Webb. Donald W,. Ph.D . Insnl Taxonomisl (iRANF.. Ph.D,. M\i oloiilSl J. [.ELAND. Bric.ham, Allison R., Pii.D . .issonatr Piofrssional Stirnti.st HiMFJ.lcK. K. B.. Ph.D., I'lanI I'alholoiiist Godfrey, (>eor(.f I... Pil D.. .4\si>tiatr Insrtt Taxonomisl Neei.y. Dan. Ph.D.. Plant I'nlholoi^ist I'nzicker. John D , Ph.D.. .isAiu lalr In.snt Taxonomisl .SciiOENEWF.iss, D, v., Ph.D.. l*liint I'alholngi.st Von;riiN. David J.. Ph.D.. Assi.stant Insnt Taxotutmist Robertson. KENNnn R . Pn D , Botanist rREWoRf.i. C^oLlN (;.. B.S.. As\4uialr SuppttrtuT Sitrnlist KNDRE.VS, Anton (i., Ph.D., Assoi tntr Botanist Bi'TCHER. Matthew K. M.S., A\Mslant Supportnr Sarnlifi IvFRsoN, I. oris R., Ph.D.. AsMstanI Botanist Gardner. Genf. M.S.. .issi.stanl Supporln-r Stirntut Sfr(.fni. |a.mf:s K... Assoanlr Suppintwr Surnlist HoEMANN. lovcE, Ph.D.. Assistant Supportit^ Siirrtltst Nfiaon. Bfttv. Assistant Supportivr Surntist McfiiFFEN. Katiirvn. M.S.. Assistant Supporln^ Sumlisi
McKnk.hi. Bh I N., M.S.. Junior ProfrssKfnal Si irntist PFRAIT. Si'ZANNF J., .issistant Supp Wfi/fi. Mark J., M S., Assistant Supporlu-r Stim ^f SECTION OF ECONOMIC ENTOMOLOGY Biii.FR. R\ndai I,,. B.S., Junutr Profrssionol Stirn:i\l BrcH)Ks. Khovlas M . B.S.. Junior Suppotlitf Surnlist Ri EsiNK. W'h I lAM (... Ph.D., Assoi intr t.ntoinohiiiisl and llrnil Hoi 1, .\i El A .\ . B.S,. Junior \ii/>/M»rlnr Sarntist Kasprowu/. jE\NiNf M., B.S.. Junuir Suppo^tnr Stirntisi .\itifbn. Jami.s K , Ph.D., hntomolufiist Kefnf, Dennis. B.S.. Supportnr Surntnn .\KMBHtiM, Kdward J.. Pii.D., h.nlomotofiist funun Ko(.AN. MARt:os. Ph.D.. hnliniuilonist Mai mboR(., Pmti I.., MS., Srnior Trt hnual .i&sislani kt'Ht.MAN, DoNAio K.. Pii.I>.. hnlomoloiiist. h.xtrnsion Bkowir. Joan 1... B.S.. Tn hnual Assistant Maduox. Josei'H v.. Ph.D.. l:ntotnoh}nnt HtMH)N. Steven. ,\I.S.. Trihnual Assistant Randei.i.. Rosr.oE. Ph.D.. UntmnototiiU. hMrnsmn Kffvin, Khomas M.. M-S.. Trthnuat .i\si\tant B()ti>E\iAN, John K,. M.S., Assminlr hnlonioh^isl KiRTN, .\nn, B.S. Trihnual .issistant Kasiman. Catherine K.. PhD., .issm intr Kniomotonisl MiiF.s, Chari.ene. Trihnual Assistant Kfi.soi. Alt.AN S.. Ph.D.. .4s\iHiittr hlntinnoloiiisl PosMiA, SiE^F. Trt hnual .issistant Irwin. Michafi. 1-... Ph.D.. Assimntr h.ntinnohmist 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 7 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 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) (as C. argillicola, also listed for Illinois are conspicuous components of the by Faxon 1890). aquatic environments of Ilhnois. Al- A doctoral dissertation submitted to though only 23 species are known to the University of Illinois in 1955, The inhabit the state (Table 1), they are Biology of the Crayfishes of Central among our largest aquatic invertebrates and Southeastern Illinois (Brown 1955), and often are present in large popu- was based on extensive collecting (at 4 10 lations. stations) in the Sangamon, Wabash, and HISTORICAL STUDIES OF Table 1. —Classification of the crayfishes ILLINOIS DECAPODS and shrimps of Illinois, primarily following Holthuis (1952) and Hobbs (1974b). Most of the historical information available on crayfishes and shrimps of^ Order Decapoda Illinois is that published by Forbes Family Palaeinonidae Genus Macrobrachiurn (1876), Rietz (1912), and Brown (1955). 1. M. ohione (Smith) of Illinois Crustacea" was, The "List Genus Palaemonetes as Forbes (1876) noted, "a first contri- 2. P. kadiakensis Rathbun bution to the knowledge of our Crusta- Family Cambaridae cea," and contained the "results of a Subfamily Cambarellinae Genus Canibarellus single season's work." Included in the Subgenus Dirigicambarus list ohione, Pal- were Macrobrachiurn 3. C. shufeldlii (Faxon) aemo7ietes kadiakensis (as P. exiUpes), Subgenus Pandicambarus Procambarus acutus (including Cam- 4. C. puer Hobbs Subfamily Cambarinae barus stygiiis), P. gracilis, Orconectes Genus Procambarus O. propinquus, O. rusticus immunis, Subgenus Girardietla (as O. placidus), O. virilis (including 5. P. gracilis (Bundy) O. wisconsinensis), and Cambarus Subgenus Ortmannicus diogenes (as C. obesus). Hagen, in his 6. P. acutus (Girard) 7. P. viaeinridis (Faxon) earlier (1870) monograph on North Subgenus Scapulicambarus the American crayfishes, had noted 8. P. clarkii (Girard) occurrence in Illinois of all of the above Genus Orconectes crayfishes except P. gracilis. Hagen 9. O. illinoiensis Brown 10. O. tmniunis (Hagen) erroneously also recorded for Illinois 11. O. indianensis (Hay) P. troglodytes, an Atlantic Coast 12. O. kentuckiensis Rhoades species. 13. O. lancifer (Hagen) A bachelor's thesis submitted to the 14. O. placidus (Hagen) University of Illinois in 1912 (Rietz 15. O. propinquus (Girard) 16. O. rusticus (Girard) 1912) and entitled Ecological Relations 17. O. stannardi Page Illinois to the of the Crawfishes of added 18. O virilis (Hagen) list of Illinois species O. indianensis, C. Genus Fallicambarus robustus (also listed for Illinois by Subgenus Creaserinus 19. F. fodiens (Cottle) Faxon 1885), C. tenebrosus (as C. bar- Genus Cambarus toni brevis), and Fallicambarus fodiens Subgenus Erebicambarus 20. (.'. tenebrosus Hay 21. C rusticijormis Rhoades This papti is published by aiilhoriiy of the Subgenus Lacunicambarus .Stale of Illinois and is a contribution of the Ser- 22. (.'. diogenes Giraid lion of Kaunistif .Surveys and Insect Identification •Subgenus Puncticamhnrus of the Illinois Natural History Survey. Dr. Lav\- 23. (',. robuslus Girard rence M. Pa^e is a Zoologist in the- .Sec tion. 335 336 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 Ohio river systems. To the growing list tered, only the early portion of the of lUinois species, including Cambar- sample was preserved, and subsequently ellus shujeldtii and O. lancijer reported collected individuals were returned in the interim by Faxon (1914), Brown to the water. added P. clarkii, O. kentuckiensis, and Specimens usually were collected by O. illinoiensis (described by Brown in dipnetting, minnow seining, or by 1956). digging them from their burrows. Statewide collections made for the Malacostraca require cover, and the present survey have added Cambarellus most successful method of collecting puer, P. viaeviridis, O. placidus, O. was to place a dip net downstream from, sta?}nardi, and Cambarus rusticiformis or (in standing water) next to, accumu- to the list of Illinois species. The list lations of stones, brush, living vegeta- of Illinois shrimps and crayfishes now tion, etc., and to kick through the stands at 23 species. material, thereby dislodging specimens into the dip net. The most successful METHODS method of collecting burrowers was to Collecting for the present survey of dig into the burrow down to the water Illinois decapods spanned a decade table, splash the water, wait for the (March 1972 to September 1982). Col- curious crayfish to investigate the dis- lections were made at 1,294 localities turbance, and grab it. This worked well (Fig. 1) in all counties of the state (Fig. in the relatively shallow burrows of 2), but were most heavily concentrated C. diogenes but less well in the deepjer in southern Illinois, where the largest burrows of P. gracilis. P. viaei'iridis, diversities of habitats and species occur. and F. fodiens. All species of aquatic Malacostraca Specimens were placed in 10-percent (Isopoda, Amphipoda, Decapoda) formalin if large, in 70-percent ethyl present at each site were collected. A alcohol if small enough to go into a plethora of taxonomic problems among 3-dram \ial. In the laboratory, speci- the isopods and amphipods prevent mens were washed in water, identified, their inclusion in this report. Lewis counted, sexed, and placed in 70- & Bowman (1981) and Lewis (1982) re- percent ethyl alcohol for permanent cently have published distributional storage in the Illinois Natural History studies on the subterranean isopods of Survey collection. Records were kept Illinois. on the presence of form I males and of females carrying eggs or young. A dot on Figure 1 indicates that one or more malacostracans were collected Measurements of crayfishes are given at that locality. Stations sampled at in millimeters of carapace length (CL). which no species were found are not Length-frequency distributions are shown on the map. Although the ab- presented for those species for which sence of malacostracans at a site can be no published growth or longevity data informative, it may indicate only that are available and of which large col- insufficient time was spent in searching. lections were made in Illinois. Keys, Old records (pre- 1972) are included on diagnoses, and descriptions are based the species distribution maps when they on Illinois populations only. Taxa are document significant changes in distri- arranged phylogenetically. except bution (e.g., for Mac robrachiuin within Orcoiifi tes where species arc ohione). A lew recently (1984) dis- arranged alphabeticalh because of our covered localities are plotted for O. present lack of understanding of inter- rusticus and O. stannardi. specific relationships. Because ecological analyses were EXTRALIMITAL SPECIES AND attempted for each species, all indi- viduals encountered usually were pre- STUDIES served; if large numbers were encoun- 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 and in Kentucky, O. bisectus and O. western Kentucky in streams draining tricuspis. It is more likely, however, directly into the Mississippi and Ohio that additions to the Illinois fauna will rivers, and they eventually may be result from human introductions, found, presumably as waifs, in Illinois. deliberate or otherwise, and could In Missouri, these are Orconectes har- originate from anywhere. risoni, O. hylas, O. luteus, O. medius, Regional publications on freshwater O. nais, O. peruncus, P. punctimanus, decapods of North America are avail- O. quadruncus, and Cambarus hubbsi, 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), southern Illinois (Fenneman 1938). California (Riegel 1959), Colorado During the Mesozoic, the seas enlarged, (Unger 1978), Florida (Hobbs 1942), and by the Cretaceous they inundated Georgia (Hobbs 1981), Indiana (Hay the Coastal Plain as far north as south- 1896; Eberly 1955), Iowa (Phillips 1980), ern Illinois. The subsequent uplifting Kansas (Williams & Leonard 1952), of the continent in the late Cretaceous Kentucky (Rhoades 1944a), Louisiana exposed much of the Coastal Plain, in- (Penn 1952, 1956, 1959; Penn & Marlow cluding the flatlands persisting today 1959), Maryland (Meredith &: Schwartz as swamps along the lower Ohio River I960), Michigan (Pearse 1910; Creaser in southern Illinois (Fenneman 1938). 1931), Nebraska (Engle 1926), New In the late Tertiary the Mississippi Jersey (Fowler 1912; Francois 1959), River ran along the western edge, and New York (Crocker 1957), Ohio (Turner the lowermost Ohio River followed the 1926; Rhoades 1944b), Oklahoma eastern edge, of Crowley's Ridge (a (Creaser & Ortenburger 1933; Reimer ridge on the Coastal Plain of south- 1969), Ontario (Crocker & Barr 1968), eastern Missouri and northeastern Pennsylvania (Ortmann 1906), Texas Arkansas). The two great rivers met near (Penn &: Hobbs 1958), West Virginia- where Helena, Arkansas, is now (Fenne- (Newcombe 1929), Wisconsin (Creaser man 1938; Fisk 1944). The Ohio River 1932), the Cumberland Plateau and was comparatively small, had its head- Cumberland Mountains (Bouchard waters in southern Indiana and central 1976), New England (Crocker 1979), the Kentucky (Wayne 1952), meandered Ozark Plateaus and Ouachita Provinces across southern Illinois through the (Williams 1954), the Southern Appa- present Cache River valley, and, after lachians and Cumberland Plateau being joined by the Cumberland and (Ortmann 1931), and Mexico (Villa- Tennessee rivers, flowed south to its lobos 1983). confluence with the Mississippi. It is Extremely useful publications on uncertain whether the Tennessee River crayfishes are Hobbs' key and check- joined the Ohio above the entrance of list to all described crayfishes of North the Ohip into the Cache lowland or America (Hobbs 1972a, 1974b). whether it flowed from its present mouth down the present course of the GEOLOGICAL HISTORY OF Ohio and Mississippi rivers and joined ILLINOIS AND the Ohio near southern Tennessee. ZOOGEOGRAPHY OF NATIVE The Mississippi River followed much of its present course in Minnesota and ILLINOIS DECAPODS Wisconsin, then cut through northern Prior to settlement by Europeans, Illinois (Willman & Frye 1970), where Illinois was covered by oak-hickory it was joined by its major tributary, forest, maple-basswood forest, and Teays River, and flowed south along the bluestem prairie. Today much of Illi- present course of the Illinois River. The nois has been cleared and developed, Teays River began along the western primarily for agriculture and urbaniza- side of the Blue Ridge Mountains of tion. Kentucky, Virginia, and West Virginia, The oldest prominent physiographic flowed northward to central Ohio, then features of Illinois are remnants of the turned west across central Indiana and Appalachian Revolution near the close Illinois through the Mahomet Valley to of the Paleozoic. The intense folding the Mississippi (Horberg 1945; Wayne and faulting during the formation of 1952; Teller 1973); or in Indiana turned the Appalachian Mountains, and the south through the present Wabash more moderate folding and tilting to Valley to the Ohio River (Fidlar 1948). the west, created the template for the It may be that an early Pleistocene ice topographic diversity present today in 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 COASTAL PLAIN the Kansan ice cap extended farther Fig. 3. —Glacial boundaries and major ptiys- south than other Pleistocene glaciers iographic features in Illinois. and terminated in Missouri, approxi- mately along the present course of the points of the moraines became drainage Missouri River. East of the Mississippi outlets. Water flowing from one lake River the Kansan ice cap was exceeded to another along the concentric mo- in its southern extent by the later Illi- raines gradually formed streams and, noian glacier, which in some areas eventually, complete drainage systems. reached the present course of the Ohio With drainage, the process of erosion River. The Illinoian leveled or buried began. As sediments were carried into in drift most of Illinois. Because it was lakes, the coarser particles (sand and the last of the glaciers, drift of the Wis- gravel) were deposited and the smaller consinan covers a large area of Illinois particles (silt and clay) were carried (Fig. 3) and its ecological impact is away. Eventually, the lakes became enormous. marshes and. later, prairies. Thick As the ice caps retreated, huge quan- layers of sand were sometimes left be- tities of drift were left behind, filling hind, such as those found today in the river channels and valleys. Mounds of Kankakee Ri\er system of Illinois and drift (moraines) sometimes reached Indiana. Dunes were subsequently heights of 30 m or more and widths of formed as the wind piled sand into several kilometers. Moraines generally hills. were concentric to one another and, Throughout the Pleistocene, an area when they formed extensive lakes, of southwestern Wisconsin and north- lacked drainage outlets. Ultimately, low western Illinois remained imglaciated. August 1985 Crayfishes and Shrimps of Illinois 341 This "Driftless Area" supports a topo- southern edges of the glaciers. Because graphic diversity, due in part to recent all of northern North America (48° and erosion, absent in the glaciated region. north) except a large part of Alaska was Interglacial periods, the Aftonian, covered by ice, the only possible non- Yarmouth, and Sangamon, were char- southern postglacial origins of the acterized by warmer and drier climates decapod fauna of Illinois are east coast similar to that of today (Braun 1950). drainages (presumably via the Sus- Outlet connecting the eastern The Wisconsinan ice cap was up to quehanna Great Lakes and the Susquehanna 3 km thick and in some areas, melting, interrupted by periodic advancement, River), the Driftless Area of north- western Illinois, the took more than 10,000 years (Clark & and upper Missouri River drainage. species Stearn 1960). New drainage systems Some may have eroded the glaciated area, and the junc- invaded from more than one refugium. tion of the Ohio and Mississippi rivers Species likely, because of their present moved progressively northward (Mat- distributions, to have invaded from the south are thes 1933; Fisk 1944). The Great Lakes M. ohione, P. kadiakensis, P. acutus, F. diogenes. were formed as water was impounded ip fodiens, and C. the former Laurentian River basin. The All but C. diogenes are sluggish-water inhabitants that probably invaded the present drainages of Illinois (Fig. 4) are discussed in detail by Forbes & Richard- low-gradient drainages of Illinois from similar low-gradient streams the son (1908) and Smith (1971). on Pleistocene glaciers that covered most former Mississippi Embayment. C. diogenes of Illinois had profound effects on the presumably moved from the distributions of organisms. The princi- Shawnee Hills and other southern areas into drainages to the north. pal zoogeographic effects were the new elimination of populations (probably P. gracilis, O. im?nunis, andO. I'irilis western drainages including entire species) and the dis- now occupy that were placement of species farther south than unglaciated, principally the upper they had occurred preglacially. Princi- Missouri, and they probably entered pal postglacial zoogeographic events Illinois from the west. The eastward were the reinvasions of glaciated regions movement of other grassland and plains and the crossing of previously insur- animals during the recent Xerotliermic mountable basin divides through the Period is discussed by Smith (1957). newly established drainage systems. O. propinquus now occurs only in The Illinoian glaciation reached glaciated areas of eastern North America within 30 km of the northern boundary but is most closely related to species of the Mississippi Embayment in south- living in streams in unglaciated south- ern Illinois (Willman & Frye 1970). A ern Ohio, eastern Kentucky, West V'ir- few species may have maintained pop- ginia, and Pennsylvania (Fitzpatrick ulations in the surviving uplands, the 1967). It and C. robustus probably in- Shawnee Hills of southern Illinois vaded Illinois from the east. C. robustus, (Fig. 3). as discussed in the species account, has since retreated from Illinois. It must have been easy for animals to invade the new drainages of the gla- O. stannardi does not now occur in ciated region of Illinois as they became areas considered to have been Pleis- connected to the existing Ohio and tocene refugia, and it may i)e auto- Mississippi rivers. Postglacial invasions chthoncjus to the Little Wabash River (cjr reinvasions) undoubtedly were system of Illinois. Autochlhonism mostly from the south; i.e., the Ozark would require it to have differentiated Plateau, Mississippi Embayment, In- from its closest relatives since the terior Lowland, Appalachian Plateau, Pleistocene. and ephemeral drainages along the 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 O. itlinoiensis M. ohione P. gracilis P. acutus C. shujeldtii O. indianensis P. kadiakensis O. propmquui O. immunis P. clarkii O. kentuckiensis O. slannardi F. jodiens P. viaeinridis O. placidus O. virilis' C. diogenes O. tancifer C. tenebrosus" C. robuslus "Also found on the Coastal Plain. .\lso found in one locality in east-central Illinois. may be divided into Coastal Plain, and becomes motionless. The male Shawnee Hills, Lowland, Glaciated- turns the female on her back, holds her area, and Statewide species groups chelae in his, positions his abdomen (Table 2). The Statewide species group over hers (still curved forward), inserts includes two burrowers (C. diogenes the ischial hooks on his pereiopods into and F. jodiens) and two sluggish-pool the coxal membranes of the pereiopods inhabitants {P. acutus and O. im- of the female (which locks the two cray- munis). fish together), and inserts the tips of Those sfjecies found on the Coastal his gonopods in the annulus ventralis Plain and in the Shawnee Hills pre- (seminal receptacle) of the female. sumably have been there for a long time Sperm travel from the reproductive and have undergone little or no post- openings on the bases of the fifth pe- glacial dispersal. Those in the glaciated reiopods along the gonopods to the regions must have invaded or re-invaded annulus ventralis, and mating is com- after glaciation (i.e., within the past pleted. A waxy sperm plug projecting 10,000-100,000 years) or must have from the annulus indicates that the speciated since then. Illinois' endemic female has recently mated. decapods, O. illtnoiensis and O. slan- Frequencies of form I males among nardi, and its near-endemics, O. ken- Illinois collections of crayfishes indicate tuckiensis and O. indianensis, occur that most matings occur between Sep- south of the boundary of the Wiscon- tember and March, although consider- sinan glaciation. All except O. slan- able interspecific variation occurs. nardi, which occurs on Illinoian glacial Species of Cambarellus have a high till, are occupants of the Shawnee Hills. incidence of form I males throughout the year. NATURAL HISTORY Sperm are carried by the female until Crayfish Life Cycle oviposition (egg laying), which in During the mating season, form I Illinois occurs mostly in March, April, males (males in which one or more of and May but may begin in some species the terminal elements on the gonopods as early as December or January. In [first pleopods] are corneous) actively preparation for oviposition the female seek receptive females by grasping other cleans the underside of her abdomen, crayfishes with their chelae. When a mostly through brushing motions of male or nonreceptive female is grasped, the fifth pereiopods. During oviposi- it resists, and the aggressive male re- tion the female turns on her back, curls leases his catch and continues search- the abdomen forward, and from glands ing. When a receptive female is grasped, on the abdomen secretes a clear, sticky it stretches the antennae, chelae, and substance (glair) which fills the area walking legs forward, curves the end of between the curled abdomen and the the abdomen ventrally and anteriorly. 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 late summer or early fall. Sexual matu- and external openings of the oviducts, rity in the male corresponds to a molt respectively. Following oviposition, to form I. The first molt following the the glair hardens and cements the fer- mating period returns the male to a tilized eggs to the pleopods. With eggs nonieproducti\e state (form II) in attached to her abdomen, a female cray- which the gonopods are without cor- fish is said to be ovigerous or "in berry" neous tips. Following copulation, (Fig. 5). females do not molt until after the Eggs are carried by the female for young permanently have left. In Illi- 2-20 weeks, depending on the water nois, crayfishes appear to survive a temperature. When the eggs hatch, the maximum of two reproductive seasons. young remain attached to the abdomen In nonburrowing Illinois crayfishes, and are carried by the female througfi all aspects of their life history occur in three instars. The first juvenile instar surface waters. Burrowing species spend has extremely large eyes, a yolk-filled variable amounts of time in surface carapace, an incomplete abdomen, and waters. Adults of P. gracilis and P. hooked chelae. First instars attach to I'iaeviridis leave their burrows and enter the female with the hooked chelae and temporary bodies of v\ater to mate. F. by a stalk linking the telson of the instar fodiens leaves its burrows to mate, and to the abdomen of the mother. After females remain out of their burrows 2-7 days first instars molt to second during the period in which they carry instars, which are more adultlike but their eggs and young. Young of these still lack a fully developed abdomen three species remain above ground in and cling to the mother's abdomen by temporary water as long as the water their chelae. After another 4-12 days, remains; as the water table recedes below another molt results in the third instar, the stnface, they burrow, .\dults of P. which closely resembles the adidt and I'laei'iridis and F. fodiens may be foimd hangs onto the mother with its chelae above ground in Illinois onl\ during and pcreiopods. It may leave the protec- periods of late winter-to-spriiig flood- tion of the mother for short periods. ing (January to May). Sexually mature Subsequent instars are free living. P. gracilis leav e bmrows on warm lainv Growth proceeds through a series of nights but otherwise seldom are found (i-10 molls during the summer, and out of their burrows, .\dults of C. dio- sexual maturity may be attained by genes leave their burrows on the banks August 1985 Crayfishes and Shrimps of Illinois 345 of streams more often than do the six stages in about 3 weeks, described species just mentioned, perhaps to in detail for Palaemonetes kadiakensis forage as well as to mate, but do so more by Broad &: Hubschman (1963). Larvae frequently during the mating and egg- and juveniles increase, and adults de- and young-carrying seasons. crease, in abundance through the sum- mer. Adults disappear in late summer- early fall, corresponding to a maximum Life Freshwater Shrimp Cycle life span of about 1 year in Palaemo- Females carry eggs attached to their netes kadiakensis and 2 years in Macro- pleopods between April and August in brachium ohione. The largest adults Illinois and between February and mature and reproduce earliest in the October farther south. The eggs hatch, year. Observations on copulation ap- and free-swimming larvae pass through 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. Fig. 6.— Lateral view of Macrobrachium ohione (after Hoitfiuis 1952). cephalic process Fig. 7.— Morpfiology of crayfish gonopods (after Hobbs 1972a). (A) Mesial (left) and lateral central projection views of terminal elements of generalized Pro- caudal process cambarus gonopod. (B) IVIesial (left) and lat- eral views of terminal elements of generalized mesial process Cambarus gonopod. (C) Generalized Orco- A nectes gonopods. central projectioi Crayfishes are most easily and accu- rately identified by using characteristics of the gonopod (first pleopod) of form I mesial process (breeding) males, and in geographic B 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 width of palm length of palm hook on ischium pleopods I 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) Fig. 10. 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) Fig. 11. 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- Fig. 12. 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 c Fig. 13. 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 Fig. 14. jjury (p. 366) 4. Terminal elements of gonopod straight (Fig. 15); margins of rostrum straight, strongly con- verging anteriorly; annulus ventralis extends Fig. 15. ./^ ventrallv (Fig. 16) Cambarrllus shufrldtii (p. ;i63) 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 Fig. 17. than basal margin of rostrum (Fig. 18) Orconectes lancijer (p. 400) 6. Rostrum shalknvly excavated; acumen much shorter than margin of rostrum 7 Fig. 18. Fig. 19. 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. 20. 1!^.. 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 Fig. 22. wFig. 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^ concavity (Fig. Cambarus deep 28) y^ PI 27 diogenes (p. 434) ^\^ 10. Margins of rostrum without large spines (Fig. 29) 11 Fig 28. 10. Margins of rostrum with large spines (Fig. 30) 15 Fig. 29. 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 Fig. 30. 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. 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. Fig. 33, 34) Procambarus viaeviridis (p. 378) Fig. 34, 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) Fig. 38. Fig. 37. 15. Margins of rostrum distinctly concave (Fig. 39) ^ 16 15. Margins of rostrum straight (Fig. 40) 18 Fig. 40. Fig. 39 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 Fig. 41 17. Dorsal (cephalic when lifted from body) surface of gonopod with prominent shoulder (Fig. 42) Orconectes ruslicus (p. 412) 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) Fig 43. ^b^^j^ ! 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- Fig 48. dianensis) (Fig. 47); 21 / Fig. 47. 21 . Mesial process of gonopod with spur on caudal surface (Fig. 48) Orconectes stnnnardi (p. 415) 21. Mesial process of gonopod without spur 22 Fig. 50. 22. Rostrum with large median carina (Fig. 49); gonopod with tips of elements barely di\erging Fig. 49. (Fig. 50); Orconectes propinquus (p. 406) / 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 Fig. 56. ^ * 1 1 ^ 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- "^'9 ^1 barus robuslus (p. 439) 354 Illinois Natural History Survey Bulletin Vol. 33, An. 4 6. Base of fixed finger of chela without deep im- pressions 7 Fig. 62 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 Fig. 63. 18 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. 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 72. 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- Fig. 74. nectes I'irilis (p. 417) 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. 17. Dorsal (cephalic when lifted from body) surface of gonopod with prominent shoulder (Fig. 76) Orconectes rusticus (p. 412) Fig. 76. 17. Dorsal surface of gonopod without prominent shoulder (Fig. 77) Orconectes placidus (p. 404) Fig. 77. 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. 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) Fig. 79 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. 80. Fig. 81. SYSTEMATIC ACCOUNTS ohione is the only species of Macro- brachium found in the Mississippi FAMILY PALAEMONIDAE River system. Both shrimps native to Illinois are members of the family Palaemonidae, Macrobrachium ohione (Smith) which is worldwide in distribution. (Fig. 82) Macrobrachium ohione, with 9-13 Palaemon Ohionis Smith 1874 teeth along the upper edge of the ros- Palaemon sallei Kingsley 1882 trum, is easily separable from Palae- Description. — Holthuis (1952) dis- monetes kadiakensis, which has only tinguished M. ohione from other 6-8 (almost always 7) teeth. M. ohione, American species of Macrobrachium which reaches 100 mm in total length by this combination of characteristics: (tip of rostrum to tip of telson), also is carpus of second legs as long as or much larger than P. kadiakensis, which longer than merus; telson with a dis- reaches a maximum length of only tinct posterior margin (rather than about 50 mm. gradually tapering toward a slender tip); second chelae of adult male equal Genus Macrobrachium Bate or nearly equal to one another in size Macrobrachium Bate 1868 (some species are bilaterally asymmet- Species of Macrobrachium, referred rical with the chela on one side much to as river shrimps, differ from species larger); no tubercles along cutting of Palaemonetes by possessing a hepatic edges of fingers of second chela of adult spine, lacking a branchiostegal spine, male; cutting edges of fingers of large and having the second pair of legs much chela of adult male with one or two longer than the first pair (in Palae- fairly large teeth proximally; eggs num- monetes the second pair is only slighth erous and small (about 0.5 mm in diam- longer than the first pair). eter); second pair of legs of adult male Although Macrobrachium contains with velvety pubescence on some or all aboiii 100 species worldwide (35 in the joints; rostrum with styliform apex (and western hemisphere), only four (A/. no teeth distally). ohione, M. acanthurus. M. carcinus, M. ohione is pale gray with light blue and M. oljersii) are found in the fresh- spots and a blue abdomen (Hedgpeth waters of the eastern United States 1947, 1949). .\duli females average (Hedgpeth 1949; Holthuis 1952). M. 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). and Ohio (Fig. 83). Coastal populations Females reach about 100 mm in length, inove into estuaries in spring (Gunter males about 70 mm (Hedgpeth 1949; 1937; Hedgpeth 1949; Reimei, Strawn, Truesdale & Mermilliod 1979). Sc Dixon 1974). Distribution and Habitat. — M. In Illinois, M. ohione has been re- ohione occupies freshwater habitats corded from Cairo, Alexander County along the Atlantic coast from Virginia (Forbes 1876); Grand Tower, Jackson to Florida, along the Gulf Coast from County (Forbes 1876); Chester, Ran- Alabama to Texas, and north in the dolph County (McCormick 1934); and Mississippi and Ohio river systems to Shawneetown, Gallatin County (Hedg- Oklahoma, Missouri, Illinois, Indiana, 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 Fig. 83. — Total distribution of Macrobrach- S. A. Forbes at Caircj (undated). The ium ohione. five collections with locality data. 358 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 (Mount Carmel, Wabash County, 1892; counted 8,000 eggs on an 84-min female Mississippi River, Grand Tower, Jack- and observed that ovigerous females son County, 1932 and 1944; Mississippi ranged from 34 to 90 mm long. Among River, Missouri River Station, Madison Illinois Natural History Survey col- County, 1944; and Mississippi River, lections of Illinois M. ohtone (all made 1 mi S Cairo, Alexander County, 1962) between May and October), ovigerous are represented on the distribution map females are present only in a collection (Fig. 84). No M. ohione were found made 13-14 May 1932. during the present survey (1972-1982). In Louisiana, Truesdale Sc Mermil- Although perhaps still present in the liod (1979) found ovigerous females state, the species obviously has under- (50-93 mm in total length) from March gone a tremendous reduction in popu- through September. Five females, rang- lation. Little information on the ecol- ing from 54 to 80 mm, were carrying ogy of the species in Illinois is available, 6,273-24,800 eggs. Eggs were ai)out but its decline presumably is attribut- 0.5 mm across, bright orange when first able to the loss of suitable habitats as laid, and brown at later stages. Huner a result of the excessive modifications (1977) found M. ohione at Port Allen, (especially channelization, impound- Louisiana, to range from 17 to 92 mm ment, and drainage of bottomland long, to average 30.6 mm (N = 7,058). lakes) of Illinois' largest rivers. and to live a maximum of 2 years. Ovi- Life History. — In the Mississippi gerous females (N = 88) ranged from 27 River at Chester, McCormick (1934) 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. Fig. 84 —Distribution of Macrobrachium occurs in Illinois. P. ohione in Illinois. Large circles = pre-1900 kadiakcnsts palu- collections, small circles = 1932-1949 collec- dosus is widespread in the eastern tions, black dot = collection made in 1962. 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 with 3-8 articles. Scaphocerite about not enter the Mississippi River system. three times as long as wide, lamella In addition to P. kadiakensis and P. overreaching distolateral tooth. paludosus, the only freshwater species Living specimens are transparent in the United States are highly localized with green eyes, red-brown antennae, spring- and cave-inhabiting forms in and many very small red-brown specks Texas (P. antrorum, P. holthuisi, and P. on the body. The internal organs are texanus) and Florida (P. cummingi) readily visible, and a bright green vege- (Strenth 1976). tation-filled intestine is often a prom- inent feature. The species reaches a 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 Fig. 86.— Appendices masculina and interna large, pigmented. Lateral antennular (left) and entire second pleopod (right) of male flagellum with rami fused for 11-23 Palaemonetes kadiakensis (after Meehean articles; free portion of shorter ramus 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. 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 Fig. 87.—Total distribution of Palaemo- netes kadiakensis. (as P. exilipes) only from the Illinois River but described it as being very locality given on the label accompany- (ommon. A specimen in the I'.S. Na- ing the type specimen is Kodiak Island. tional Museum of Natural History Alaska, it is doubtful that the species documents its former occurrence at ever occurred in Alaska (Holthuis 1952). Kankakee. Kankakee County (pre-1898. P. kadiakensis now occurs through- exact date unknown), a localitv well out the southern one-fourth of Illinois, outside the present range. in backwaters along the Mississippi P. kadiakensis almost always is asso- River, in backwaters of the Illinois ciated with, and is most abundant in. River at least as far north as Bureau li\ing a(]ualic \egetation. Its reduction County, and in Wolf Lake (Lake Mich- in distribution and abundance in Il- igan drainage) in Cook County (Fig. linois probably is attributable to in- 88). It abounds in swamps (Fig. 89) creased turbidiiv and .sedimentation and swamplike streams on the Coastal and the resultant loss of vegetation. Plain of Illinois and is common in Beds of emergent and submerged \ege- standing- and sluggish-watei habitats t.iiion once were common throughout as far north as Salem. It is uncommon Illinois but now are rarely encountered. in the Illinois River and in Wolf Lake. 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- 1.0 mm in diameter. The egg mass con- tions of Illinois fishes (Smith 1979). stituted an average of 11.2 percent of Life History. — Reproduction in the weight of the female (without the Illinois, as indicated by the presence of eggs) in 15 Illinois females (Table 4). ovigerous females, occurs from April to Females may produce more thair one August (Table 3). In Louisiana the brood per season (Broad &: Hubschman reproductive period extends from Feb- 1963; Nielsen & Reynolds 1977). The ruary to October (Meehean 1936; White incubation period lasts 24-28 days at 1949), and in central Missouri (where 18.5°-24°C. Detailed accounts of lar- the species is introduced) from mid-May to August (Nielsen k Reynolds 1977). Table 3.— Illinois collections of Palae- In Illinois collections, ovigerous monetes kadiakensis. 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 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- southern Illinois to southwestern Ala- cambarus, the telson always is divided bama and eastern Texas (Fig. 91). In in North American cambarids. Illinois, C. shufeldtii is known only from Alexander, Jackson, Massac, Genus Cambarellus Ortmann Pulaski, and Union counties in both Cambarellus Ortmann 1905 the Mississippi and Ohio basins. Most Gonopod with three terminal ele- localities are on the former Mississippi ments; central projection not large and Embayment, but the species also as- bladelike. Ischia of second and third cends the lower M ississippi River Valley pereiopods of male with hooks. and the lower Big Muddy River to about Species of Cambarellus are distrib- Sand Ridge in Jackson County (Fig. 92). uted throughout the southern United C. shufeldtii was first collected in Il- States and Mexico, generally in swamps linois at Cairo by Robert Kennicott and other standing- or sluggish-water (Faxon 1914). Brown (1955) found the habitats. They are "dwarf" crayfishes, species in five sloughs and ditches in reaching a maximum total length of Alexander and Massac counties. The 28 about 30 mm. Two of the 17 recognized collections made in the present survey species of Cambarellus occur in south- were from cypress swamps, sloughs, and ern Illinois. Fitzpatrick (1983) revised backwaters. C. shufeldtii is especially Cambarellus, proposing the recogni- common in the heavily vegetated LaRue tion of three subgenera. Swamp-Wolf Lake complex in Union County. C. shufeldtii and C. puer have Cambarellus shufeldtii (Faxon) not been found together in Illinois and (Fig. 90) apparently compete for suitable habi- Cambarus Shufeldtii Faxon 1884 tats. In Gulf Coast streams C. shufeldtii Description. — Rostrum broad, flat, is supplanting C. puer (see C. puer often deflecting downward anteriorly; account). margins straight, strongly converging C. shufeldtii was probably more anteriorly, ending in large spines; common and widespread in extreme acumen moderately large. Carapace southern Illinois prior to the extensive compressed, with cervical spines; sub- clearing of swamps and river flood- orbital margin angular. Areola wide, plains. Brown (1955) mentioned an narrowest part about 19-24 percent of earlier collection of a "dwarf crayfish" length. Chela narrow and long, smooth; near Shawneetown (Gallatin County) palm without tubercles. Form I gono- but was unable to document the occur- pod terminates in three elements, all rence of Cambarellus that far northeast. distally directed (not curved): sclerotized In view of the occurrence, or former central projection, nonsclerotized occurrence, of several swamp-inhabit- mesial process, and nonsclerotized ing fishes (Fundulus dispar, Elassoma caudal process. Dorsal color variable, zonatum, Lepomis punclatus, and L. from rust red to light brown with two symmetricus [Smith 1979]) that far dark brown stripes or rows of spots on northeast, or even into the lowlands either side of the areola extending down of the lower Wabash River drainage, the abdomen. the presence of C. shufeldtii or C. C. shufeldtii appears to have no ob- puer in the same area is possible. viously close relatives, being the only In Louisiana, C. shufeldtii lives in species in the genus with straight ter- clear, shallow (less than 40 cm deep), minal elements on the gonopod and the permanent, sun-exposed, mud-bot- only member of the subgenus Dirigi- tomed, vegetated bodies of water (Penn cambarus (Fitzpatrick 1983). 1950). C. shufeldtii characteristically Distribution and Habitat. — C. does not burrow but may survive peri- shufeldtii occupies bodies of sluggish ods of drought in subterranean (ells water, mostly on the Coastal Plain, from (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: ventralis; C, annulus 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 Fig. 92.— Distribution of Cambarellus shu- desiccation. to prevent feldtii in Illinois; black dots = 1972-1982 col- Life History. — Form I males are lections. present in Illinois collections (Table 5) during most months and peak in occur- CL, collected in February and March rence in December, February, and were carrying 47-99 (mean, 80.1) eggs March. In Louisiana, Penn (1942, 1950) averaging 1.1 mm in diameter. The re- found form I males in all months except lationship between numbers of eggs (N) September, with peaks in January, and carapace length (CL) is N = —539.0 February, and July. Presumably, + 558.9 log CL, r = 0.86. Four females, mating in C. shufeldtii may occur at 12.4-13.5 mm CL, collected in April anytime but peaks in winter in Illinois 1974 were carrying 15-74 (mean, 44.8) and in winter and summer in Louisiana. young. The decline from an average of Females carrying eggs were found in 80.1 eggs to an average of 44.8 young Illinois from February to May, and suggests a 44 percent mortality during peaked in occurrence in March (Table egg and early instar stages. Penn (1942) 5); females carrying young have been found the number of eggs or young on found in Illinois in April, June (Table a female to increase with the carapace 5), and July (Brown 1955). In Louisiana, length of the female, with females 8.5 females with eggs or young are jjiesenl mm averaging 27 offsjiring and those in every month, with strong peaks in 1 1 mm averaging 54 offsjjring. \n oxer- January-Mardi and June-July (Penn all average of 34 offspring per female 1942, 1950; Lowe 1961), again suggest- {N = 100) in Louisiana (Penn 1942) is ing two peaks of repiodut ti\i' activity about one-half the average number of in the southern part of the range. ()ns|)riiig (69.2) [jer female (;V = 13) in Nine Illinois females, ll.tJ-14.2 mm 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 (Fig. 97), effluent streams, and lowland anteriorly, ending in large spines; areas which probably were swamps acumen moderately large. Carapace prior to being cleared. \\\ but one col- compressed, with cervical spines; sub- lection from a roadside ditch were made orbital margin angular. Areola wide, in permanent water bodies, and most narrowest part about 25 percent of specimens were found among living or length. Chela narrow and long, smooth; dead vegetation. C. puer may be more palm without tubercles. Form I gono- widespread on the Coastal Plain of Il- pod terminates in three elements: sclero- linois than present records indicate and tized central projection, nonsclerotized almost certainly was more widespread mesial process, and nonsclerotized and common prior to the drainage of caudal process, all curved at sharp angle swamps and lowlands. (about 45°-90°) to principal axis of C. puer and C. shujeldtii were col- gonopod. Color as in C. shujeldtii. lected at 23 sites in Illinois, mostly The closest relatives of C. puer are C. in Coastal Plain swamps, but never schmitti and C. lesliei (see Fitzpatrick were collected together. Competitive &: Laning 1976), two occupants of the exclusion among species of Cambarel- Gulf Coastal Plain from Mississippi lus has been documented in Gulf Coast to Florida. C. puer, C. schmitti. C. areas (Penn & Fitzpatrick 1962. 1963) lesliei, C. diminutus, C. blacki, C. and apparently is operating in Illinois. ninae, and C. texanus all are members C. puer and C. shujeldtii seem unable of the subgenus Pandicambarus (Fitz- to coexist and, at least along the Gulf patrick 1983). Coast, C. shujeldtii has supplanted Variation in morphological charac- C. puer at several localities within teristics of C. puer was studied by historic times (Penn &: Fitzpatrick 1962. Chambers, Payne, & Kennedy (1979). 1963). C. shujeldtii was found experi- Three populations were separable from mentally to be dominant o\er C puer one another but not given taxonomic (i.e., C. puer retreats following contact status. Illinois populations are part with C. shujeldtii) (Penn &: Fitzpatrick of a larger population extending along 1963), suggesting that in nature C. the Mississippi River and across south- shujeldtii agressi\ely displaces C. puer ern Louisiana. In general, individuals from certain habitats. in the east are larger than those in the Life History. — Louisiana data sug- west; females are larger and have gest IWC3 ieproducti\e periods (winter shorter, broader chelae and shorter and summer) (Penn 1950; Black 1966). rostra than males. do although form I males can be found Distribution and Habitat. C. puer in all months (Black 1966). Illinois ranges from southern Illinois and data are seasonally incomplete (Table southeastern Missouri to the Gulf Coast 6), but two reproductive peric:)ds. some- of Louisiana and the lower Colorado what later (i.e.. late winter-spring and River drainage of Texas (Fig. 95). All fall) than in Louisiana, may also occur populations are on the Coastal Plain. in Illinois. In Louisiana, form I males In Louisiana, C. puer occurs most peak in occurrence from December to frequently in shallow (less than 40 cm March (abc^ut 60-95 percent of all males) deep), clear, permanent, sun-exposed, and from July to .\ugust (about 45-75 mud-bottomed, vegetated water bodies percent) (Penn 1950: Black 1966). In (Penn 1950). the available Illinois collections (Fable C. puer was first found in Illinois in 6), form I iiiak's are present February- 1973 (Page & Burr 1973) and now is May and in October. In Louisiana, form known fr(jm seven sites in the Missis- I C. puer males measure 7.2-12.0 mm sippi and Ohio basins in I'nion, John- CL and weigh 75.4-412.1 mg (Black son, and Alexander (ouniies (Fig. 96). 1966). Fhirteen or 14 molts are required In Illinois, it occupies cypress swamps to reach sexual maturity, with a mean August 1985 Crayfishes and Shrimps of Illinois 369 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- Fig. 96,— Distribution of Cambarellus puer niinuiicus (aculus and iiian'iridis). and in Illinois; black dots = 1972-1982 collections. .S( afj u lit a )n bams (da rk li). 370 Illinois Natural History Survey Bulletin Vol. 33. An. 4 Procambarus gracilis (Bundy) Areola extremely narrow, almost linear. (Fig. 98) Chela large, hea\ily punctate; palm Catnbarus gracilis Bundy 1876 with one row or large tubercles on Description. — Rostrum broad, deejjly mesial maigin; dactyl with shallow exca\aied. with short acumen, no concavity on basal half of opposable median carina; margins barely converge margin. Form I gonopod terminates in anteriorly, without tubercles or spines. four short elements: a sclerotized, curved Carapace compressed, lacking cervical central projection; a curved mesial spines; suborbital margin angular. 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 Fig, 100. — Distribution of Procambarus May. Brown (1955) found form I males gracilis in Illinois. Open circle = 1922 record in Illinois in August and October and for the Kankakee River; black dots = 1972-1982 females with young attached in March, collections. April, and October. Brown's largest form I male was 39 mm CL, and his noted that "females with young at- largest female was 47 mm CL. Williams tached are taken in Missouri as late as & Leonard (1952) found young attached Octobei." to many of the females collected in Among all specimens collected in the early spring in Kansas. Greaser (1932) 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. Males ( Form I) _i 5 -1 ca rH Tif Vl.n Females 5 - 1 nn„^^ 5 10 15 Fig. 102.— Capped entrance to burrow of Procambarus gracilis in a field in Des Plaines, Cook County, Illinois, 15 June 1972. — 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. ) An. 4 Natural History Survey Bulletin Vol. 33, 376 Illinois the P. blan- P. acutus is a member of Procambarus acutus (Girard) subgenus dingii species group of the (Fig. 104) Ortmannicus (Hobbs 1962, 1972b). Two 1852 Cambarus acutus Girard subspecies of P. acutus are recognized. stygius Bundy 1876 occupies San Luis Cambarus P. a. cuevachicae broad pos- with Description.— Rostrum Potosi and Puebla and intergrades converges anteriorly, Mexico and teriorly, strongly P. a. acutus in northern carina, marginal the rest excavated, no median Texas. P. a. acutus occupies small to absent species (Hobbs spines or tubercles of the range of the individuals). Carapace ( 1981 (largest on small 1972a), as described below. Hobbs spines present; regionally compressed, cervical noted the existence of se\eral angular. Areola acutus^nd suborbital margin restricted "variants" of P. a. 5-8 per- narrow, narrowest part about suggested that some may deserve taxo- and narrow, cent of length. Chela long nomic recognition. row of heavily piinctate; palm with Cambarus stygius was described bv mesial margin, one specimens large tubercles on Bundv (in Forbes 1876) from on dorsal and shore or two adjacent rows found, following a storm, on the Form I gonopod Wisconsin. ventral surfaces. of Lake Michigan at Racine, all more terminates in four elements, (Bundv 1882) and listed by Forbes (18/6) caudodistally and vari- Creaser (1932) or less directed as an Illinois species. of setae origi- syno^ ously obscured by a mass declared Cambarus stygius to be a knob: a relatively acutus (- nating on the caudal nym of Cambarus blandmgii projection, an large sclerotized central Cambarus acutus Girard). process, unsclerotized slender mesial Distribution and Habitat.— P. acutus cephalic process, and a east it occurs a sclerotized has a disjunct range. In the process. Dorsal color Maine sclerotized caudal along the Atlantic Slope from to dark red; largest in- Rner sys- pinkish brown to central Georgia (Altamaha with a black rectangle and dividuals dark red tem) and to the west (Great Lakes segment. from on each abdominal Mississippi Ri\er basins) it ranges Indiana southern Michigan and western and west to southeastern Minnesota Oklahoma, south to Alabama, western Mexico and along the Gulf slope into (Fig. 105). P acutus has been counted among (Hagen the Illinois fauna since 1870 stated that it 1870), and Forbes (1876) was very common in ceniral Illinois. Cam- Rietz (1912) found the species (as distrib- barus blaminigii) to be widely docu- uted and generally abundant and Illinois mented its occurrence in ;}2 it 36 counties. Brown (1955) found m counties drained by the Sangamon Wabash, and Ohio river systems and in the reported it especiallv common Sangamon system. Populations m Il- to as linois were generally referred P blaniiingii until Hobbs (1962) re- blanclmgn) stricted the range of P. a. to North and South Carolina. most, Presently P. acutus occupies Fig. 105.—Total distribution of Procambarus state, but perhaps all, drainages of the acutus. August 1985 Crayfishes and Shrimps of Illinois 377 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 Fig. 106 —Distribution of Procambarus explain the extremely small number acuius in Illinois; black dots = 1972-1982 col- lections. of ovigerous females found in Illinois and elsewhere (see below). In the Reel- records are absent from the Spoon and foot Lake area, Hobbs & Maichand other small rivers in western Illinois and (1943) found burrows of P. acutus to from the Pecatonica and other streams be shallow excavations, usually under in northwestern Illinois (Fig. 106). It is logs and consisting of a single passage- most common in the sluggish streams 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 largest Illinois specimen is a 54.1-mm along the margin of a drainage ditcfi CL female collected on 20 February in Iowa (Phiillips 1980). 1974 in the Maeystown Creek system, Life History. — In the present survey Monroe County. The largest male is a of Illinois, 212 collections of P. acutus 54.0 mm-CL form I collected in 1974 were made. Form I males were found in the Mississippi River, Whiteside every month except October and No- County. Hobbs (1981) recorded a 58.9- vember (Table 8). They peaked in rela- mm CL form I male from Georgia. The tive abundance in June-September and, smallest form I male from Illinois is secondarily, in January. Cralley (1932) 30.8 mm CL and was collected in the found form I males in Illinois in June Mississippi River, Randolph County, and July, and Brown (1955) from April on 24 March 1975. to July and in September. Form I males (Faxon) have been found in June and July in Procambarus viaeviridis (Fig. 107) western Tennessee (Hobbs &: Marchand 1943); in August and September in the Cambarus viae-vnidis Faxon 1914 Ozark-Quachita region (Williams Description. — Rostrum flat, deflect- 1954); from February to July and in ing downward anteriorly; margins con- September in Texas (Penn & Hobbs verge anteriorly to short acumen, with- 1958); from May to August in Iowa out spines or tubercles; no median (Phillips 1980); in April, August, and carina. Carapace compressed, lacking "the fall" in Georgia; in March, April, cervical spines; suborbital margin June, July, and September in Alabama; barely angular. Areola narrow, narrow- in April and May in Florida; and in all est part about 4-7 percent of length. months except May in South Carolina Chela long and narrow, heavily punc- (Hobbs 1981). tate; palm with row of 7-9 long tuber- No females were found carrying eggs, cles on mesial margin. 2-3 adjacent and only one (49.3 mm CL), collected rows of smaller tubercles. Form I gono- in December, was carrying young {N = pod terminates in four short elements: 30). In other Illinois surveys, Rietz a sclerotized mesial process, a sclero- (1912) reported a female collected with tized central projection, a cephalic young in April, Cralley (1932) reported process, and a large setae-covered ovigerous females collected in July, caudal knob on cephalic surface. Adults and Brown (1955) found a female with are deep rust red dorsally with the ab- eggs in August and one with young domen somewhat lighter in color than attached in April. All females with the carapace. eggs and young found by Cralley and P. viaeviridis is a member of the P. Brown were taken from burrows. blandingii species group of the sub- Few females with eggs or young have genus Ortmanniciis (Hobbs 1962, been collected elsewhere, probably 1972b). because of their sequestering behavior Distribution and Habitat. — P. viae- (Penn 1956; Hobbs 1981). In\he south- viridis occupies standing water bodies eastern United States, no females with and sluggish streams on the former eggs and only one female with young Mississippi Embayment and Gulf (from Alabama in September) have been Coastal Plain from southern Illinois collected (Hobbs 1981). Turner (1926) south to northern Louisiana and cen- reported eggs and young on P. aculus tral Alabama (Fig. 108). in Ohio and Indiana in March, July, P. viaeviridis is restricted in Illinois and September, and Penn (1956) found to cypress swamps and floodplains females with young attached in October along sluggish streams in the extreme and January in Louisiana. southern part of the state (Fig. 109). In the present survey males slightly First reported for Illinois by Page and outnumbered females, 523 to 513. The 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 Vol. 33, An. 4 1 Fig. 108. —Total distribution of Procambarus viaeviridis. from localities in the Ohio River drain- age in Alexander, Johnson, Massac, Fig. 109 —Distribution of Procambarus Pope, and Pulaski counties. The largest viaeviridis in Illinois: black dots = 1972-1982 populations are in cypress swamps collections. {especially Heron Pond-Little Black Slough and Bell Pond) in Johnson During this period, water levels in County. southern Illinois are generally high, Life History. — Other than brief and P. I'iaei'iridis can be found on comments on its affinity for sluggish flooded floodplains, in some roadside and standing bodies of water, nothing ditches, and in swamps. Periodically appears to have been published on the flooded areas appear to be the preferred ecological characteristics of P. inaeviri- habitat of P. I'iacviridis, and when they dis. During the present study, 16 collec- dry, (he crayfish burrows underground. tions of the species were made, all from In April, specimens were dug from bur- December through May (Table 9). 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. reveal that the species may be found in 110). In December and January, 1975- permanent swamps even during this 76, crayfish were active in water 1°C period. under a cover of ice. The usual absence Form I males were present in collec- of floodwater in southern Illinois from tions made in January and May (Table June through November accounts for 9). Among the four form I males col- the lack of Illinois collections during lected, the smallest was 24.1 mm CX, this period (Table 9); howe\er, more and the largest was 29.9 mm C^L. I'he extensive effort presumably would largest Illinois specimen of P. viaeviri- dis is a 32.3-mm CL female collected in Heron Pond on 30 March 1973. No Males ( = Form I) 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. mm CL Procambaru.s ciarkii (C»iiard) (Fig. 112) Fig. 111.— Size-frequency distribution of Procambarus viaeviridis collected in Bell Pond, Ciimhariis Clarkii Ciiard 1852 Johnson County, Illinois, 19 May 1973. 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, Florida panhandle to northern Mexico terminating in small to large spines; and up the former Mississippi Embay- acumen long; no median carina. Cara- ment to southern Illinois (Fig. 113). It pace compressed, with cervical spines; is the crayfish commonly harvested for suborbital margin angular. Areola ex- human consumption in the eastern tremely narrow, sometimes obliterated United States. at midlength. Chela long and narrow, In Illinois, P. clarkii occurs in punctate; palm with row of 6-8 tu- swamps, vegetated ponds, and stream bercles on mesial margin, 2-3 adjacent pools in the extreme southern part of rows of smaller tubercles. Form I gono- the state (Pope, Johnson, Massac, pod with a large angular shoulder on Union, Pulaski, and Alexander coun- cephalic surface, a large lobiform scle- ties) (Fig. 114). Most localities are in rotized cephalic process, small mesial the Ohio River drainage, although process, and small sclerotized central some are in the Mississippi River drain- projection. Light to dark red dorsally age. It remains common, especially in and laterally; white ventrally with blue Mermel Swamp (Fig. 89), Bell Pond stripe along middle of abdomen. (Fig. 97), LaRue Swamp, Horseshoe P. clarkii is a member of the subgenus Lake, and parts of the Cache River sys- ScapuUcarnbarus (Hobbs 1972b), an tem but undoubtedly was more com- assemblage of six species confined to mon prior to the drainage of much of the Atlantic and Gulf Coastal Plain of the southern Illinois wetlands. P. clarkii the United States and Mexico (Hobbs is taken often with P. acutus but is more & Grubbs 1982). common than P. acutus in the cypress- Distribution and Habitat. —P. clarkii is an abundant inhabitant of standing water bodies and slowly flowing streams on the Gulf Coastal Plain from the Fig. 113.—Total distribution of Procambarus Fig. 1 14.— Distribution of Procambarus clar- clarkii. The southwestern limit of its range is kii in Illinois; black dots = 1972-1982 collec- unknown. 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 row of 8-9 tubercles on mesial with young attached in Illinois on 6 margin; 1-2 adjacent rows of tubercles September 1954; all were dug from bur- on dorsal surface. Form I gonopod ter- rows. Penn (1943) found a female only minates in two short, straight elements: 30.5 mm CL carrying young in Loui- a sclerotized central projection and an siana. unsclerotized mesial process. Dorsally The largest Illinois specimen is a light to dull red brown; fingers of chelae 60.7-mm CL female collected in Horse- with red tips, subdistal black bands. shoe Lake, Alexander County, on 20 O. illinoiensis is a member of the June 1973; the largest male, from the Propinquus group of Orconectes (Fitz- same collection, is a 51.7 inm CL patrick 1967). The closest relative in form I. Brown (1955) recorded a 54- Illinois is O. propinquus. mm CL form I male from Illinois. Distribution and Habitat. O. il- The size-frequency distribution of 116 linoiensis is endemic to southern Il- individuals collected in the Bay Creek linois. It was recognized first by Brown system on 19 May 1973 suggests that (1955) as a distinct species and formally individuals of both sexes live a maxi- described by him in 1956. During the mum of about 2 years (Fig. 115), al- present survey it was found to occupy though the largest female mentioned rocky riffles and pools of streams in the above, at 60.7 mm, may have been 3 Ohio and Big Muddy (Mississippi River years old. drainage) systems of Hardin, Pope. Johnson, Massac, Pulaski, Alexander. Genus Orconectes Cope Union, Williamson, and Jackson counties (Fig. 117). Most of the known 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 Fig. 117— Distribution of Orconectes illi- punctate; fingers often very long; palm noiensis: blacl< dots = 1972-1982 collections. August 1985 Crayfishes and Shrimps of Illinois 387 localities (Fig. 118) are in the Shawnee tion of the species today probably ap- Hills, but a few populations occupy proximates its prehistoric distributicjn. rocky streams on the Coastal Plain and Many populations are large, especially in the region just north of the Shawnee in upland streams in Pope County. Hills. Some populations may have been Brown (1956) recorded a collection eliminated or reduced in recent decades of O. illinoiensis from Cypress Ditch through stream desiccation or other near Junction in Gallatin County. This recent perturbations, but the distribu- 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 immunis. A, dorsal view r Ji?V^°~?'?°"l'"®^ of carapace: B. dorsal view of right chela- "'" °' '"^"^ ^°"°P°' °' ' "^^'^^ ^' '^'«-' -«- °' gonop'od oj rormTmalortn 1 male. F, lateralLtPr«, ""'f eT view of gonopod of form II male. — 390 Illinois Natural History Survey Bulletin Vol. 33, An. 4 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. 121. -Total distribution of Orconectes immunis. 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 Fig. 122.— Distribution of Orconectes im- considered sufficiently distinct to war- munis In Illinois; black dots = 1972-1982 col- rant laxonomic recognition (see syn- 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- O. immunis lives in slowly flowing burger 1933) is far south of other popu- pools, usually with soft bottoms, of lations and may have been introducted. headwaters, creeks, and srnall to large The species occurs throughout Illi- rivers (Fig. 123). It also is found in nois except in the unglaciated north- standing-water bodies but, at least in west (Fig. 122). It appears to be rare Illinois (and in the Reelfoot Lake region in extreme western Illinois, although of Tennessee, Hobbs & Marchand 1943), Rietz (1912) found it in several western is primarily a stream species. It fre- counties (Fulton, Hancock, Henderson, quents intermittent streams and tem- and Mercer) for which recent records are porary ponds, burrowing down to the unavailable. Elsewhere in the state it is water table as drying occurs. O. im- generally common, and local popula- munis can be found under rocks in dry- tions sometimes consist of thousands ing streams, and burrows are con- of individuals. It undoubtedly was even structed as a means of escaping des- more common and widespread when iccation. Burrows usually are simple Illinois had large expanses of undrained structures, although they may have prairies. In fact, Forbes (1876) referred chimneys and occasionally are relatively to O. immunis as "the commonest elaborate. Brown (1955) described the species of central Illinois," and "es- burrows as up to 2 feet deep and ending pecially frequent in the muddy ponds of in an enlarged pocket below the water the prairies, where it may be drawn by table. the hundreds with a small seine." 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 Fig. 125.—Total distribution of Orconectes 1941; Crocker 1957; Crocker &Barr 1968) indianensis. and from April through June in Iowa spines; small (Caldwell Xc Bovbjeig 1969; Phillips median carina. Carapace 1980). flattened dorsoventrally. with large Seven Illinois females collected dur- cervical spines: suborbital margin ing the present study (21.8-35.9 mm barely angular. Areola moderately CL) were carrying 102-285 (mean, wide, narrowest part 15-20 percent of 168.9)eggs. Eggs averaged about 1.7mm length. Chela large, hea\ily punctate; in diameter. Three females (26.6-35.5 palm with row of 7-9 tubercles on 2-3 mm CL) carried 58-184 (mean, 120.3) mesial margin. adjacent rows of young. smaller tubercles on dorsal surface. The largest individual encountered Form I gonopod terminates in two during the present survey was a 41.9- short, distally tapering and strongly di\erging mm CL form II male collected in Ben- elements: a sclerotized cen- nett Creek, Crawford County, on 30 tral projection and an imsclerotized April 1973. Brown (1955) collected a mesial process. Overall dark brown 46-mm CL female from Illinois, and dorsally, except thorax yellow brown; Crocker (1957) recorded a 49-inm CL fingers of chelae with red tips and sub- female from New York. The ratio distal black bands. among all Illinois specimens collected O. indianensis is most similar, and was 505 males: 513 females. perhaps most closely related, to O. wrighli. a species known only from Hardin Countv, Tennessee (Hobbs Orconectes indianensis (Hay) 1974b). (Fig. 124) Distribution and Habitat. — O. in- Cainbarns itidianensis Hay 1896 dianensis lives in rocky riffles and pools Description. — Rostrum exca\ated, of small to medium-sized streams in the slightly deflet ted downward anteriorly, Wabash Ri\er drainage of southwestern with long acumen: margins con\erge Indiana and in the .Saline Ri\er and slightly anteriorly, ending in small 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 Fig. 126.— Distribution of Orconectes indi- localities) were based on misidentifica- anensis in Illinois. Open circles = pre-1912 records (Rietz 1912); black dots = 1972-1982 tions of the then undescribed O. illi- collections. noiensis and O. stanndardi. Rietz's record for Grand Pierre Creek, Pope County, and two records for Bay Creek, Saline River and, more interestingly, Pope and Johnson counties were surely for the North Fork Saline River must based on O. illinoiensis. Her record have been for O. indianensis. No simi- for Skillet Fork, Wayne County, was lar species occurs or ever has been re- probably based on O. stannardi. The corded for the Saline system. The North Little Fox River, White County, record Fork records are interesting because may have been based on O. propinquus. O. indianensis no longer is found there. However, her records for South Fork 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 averaging 15.2 Males Form I) year-old males mm (N = 25), 1 -year-old females averaging I^ 13.4mm(N = 22),and2-year-olds(.V = 4) ranging from 25 to 33 mm CL. Of the of O. indianensis col- [0 Femal es ( with eggs) 219 specimens lected during the present survey, 104 „ were males and 115 were females. 10 20 30 40 mm CL Orconectes kentuckiensis Rhoades (Fig. 128) Fig. 127.—Size-frequency distribution of Orconectes indianensis collected in Honey Orconectes kentuckiensis Rhoades 1944 Creek, Hardin County, on 23 Marcfi 1973. Description. — Rostrum excavated, deflected downward anteriorly, with barely .Saline are badly polluted. They suffer long acumen; margins converge tubercles or small from strip-mine and oil-field pollution, anteriorly, ending in carina. siltation, and desiccation (Smith 1971) spines; lacks median Carapace with large and are among the most damaged and flattened dorsoventrally, abused streams in Illinois. In many cervical spines; suborbital margin Areola wide, narrowest areas of these streams little aquatic life barely angular. length. of any kind can be found. part about 18 percent of Chela large, hea\ily punctate; palm with row Life History. — Apparently the only of 7-9 tubercles on mesial margin; 1-2 ecological information on O. indianen- adjacent rows of smaller tubercles on sis is that presented by Brown (1955) and dorsal surface. Form I gonopod termi- that from the present survey. nates in a short, thick, distally directed, Brown (1955) found no ovigerous sclerotized central projection, and a females and collected form I males only short, caudodistally directed, unsclero- in September and November. In the tized mesial process. Dorsally dark present study, collections were made brown (with yellow brown thorax) to from March through October (except red brown; fingers of chelae with red June), and form I males were present tips, subdistal black bands. in March, August, September, and The gonopod of O. kentuckiensis October (Table 13). The smallest form most closely resembles that of O. sloanii I male was 17.7 mm CL, and the largest in southern Indiana and Ohio and O. was 33.4 mm CL. hamsonii in southeastern Missouri. I Ovigerous females were found in Although the relationships of these March and April (Table 13). One hun- species ha\e not been in\estigated. dred twenty-one eggs were counted O. sloanii and O. harrisonii probably on a 26.3-mm CL female, 148 eggs on a are the closest relatives of O. kentuck- 27.7-mm CL female, and 178 eggs on a iensis. 32.2-mm CL female. Eggs averaged Distribution and Habitat. O. ken- about 1.8 mm in diameter. The only tuckiensis occurs only in small streams female with young attached was 35.0 in a small area of southeastern Illinois mm CL and was collected on 19 May and northwestern (Crittenden. Li\ing- 1973. Attached to her were 132 young ston, and I'nion counties) Kentucky and 21 unhatched eggs. (Fig. 129). In Illinois, it occurs only The largest Illinois specimens ex- in Big, Hosick, and Peters creeks, three amined were the above-mentioned 35.0- spring-fed, rocky, direct tributaries of mm female and 33.4-mm form I male. the Ohio River in Hardin County (Fig. The size-frequency distribution of 51 130). Its distribution in the headwaters specimens collected in Honey Creek. of Big Creek (Fig. 131 ). by far the largest Hardin County, on 23 March 1973 (Fig. of the three streams, complements the 127) suggests a 2-year life span, with 1- downstream distributit)n of O. plandus 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 Fig. 130. — Distribution of Orconectes ken- tuckiensis in Illinois; black dots = 1972-1982 i Brown (1955). O. kentuckiensis prob- collections. ably was never more widespread in Il- linois than it is today. O. kentuckiensis is most common in lations of brush over mud. This species shallow, rocky pools of small streams does not burrow but may bury itself in (Boyd & Page 1978), where large rocks loose gravel 2-4 cm below the surface are used as cover, although Rhoades of a dry stream bed (Boyd k Page 1978). (1944a) collected specimens in accumu- Life History. —The life history of O. { 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 dorsoventrally, with large cervical by Boyd & Page (1978). The species spines; suborbital margin angular. was active at temperatures of 6°-32°C Areola narrow, obliterated anteriorly. and dissolved oxygen levels of 3.5-14.4 Chela long and narrow, setose, without ppm. Most individuals were found in tubercles. Form I gonopod terminates water 5-61 cm deep. The species lived in two short, spatulate elements: a a maximum of 2+ years, had a 1:1 sex sclerotized central projection and an ratio, fed on vascular plants and arth- unsclerotized mesial process. Mottled ropods (amphipods, isopods, crayfishes, overall in shades of red, brown, and caddisflies, and midges), and was preyed green; fingers of chelae without bright upon by fishes. red tips and without black bands sub- Form I males (15.9-37.5 mm CL) were distally. present from July through April and O. lancifer is a highly distinctive peaked in occurrence in November, crayfish with no obviously close rela- December, and February; females carry- tives. ing eggs or young were present in — March, April, May, and July (Table 14) Distribution and Habitat. O. lan- and measured 14.8-32.8 mm CL. Fif- cifer is an inhabitant of the Gulf Coastal teen females were carrying 49-248 Plain and is known to occur in Louisi- (mean, 110) eggs. Eggs were spherical ana, Mississippi, Tennessee, eastern southern and averaged 1.9 mm in diameter. The Texas, Arkansas, and extreme relationship between the number of Illinois (Fig. 133). It lives in deep water eggs (N) and the female's weight in of oxbows, bayous, and large streams grams (W) was N = 49.809 + 25.310 VV, (Black 1972) and, as a result is rather collections. r = 0.807, and that between N and the poorly represented in female's carapace length (L) was N = O. lancifer has been found at only -0.43611 + 1.8516 log L, r = 0.859. two localities in Illinois. Robert Kenni- Counts of ovarian eggs in 15 females cott, an early naturalist, collected a were 59-249 and averaged 109. Nimi- female at Cairo in the middle 1800's bers of eggs, ovarian and abdominal, (Faxon 1914). and Page & Burr (1973) increased significantly with carapace recorded a collection of eight individ- length. The growth of first-year cray- uals from Horseshoe Lake. Both local- fish occurred during a period of 4.5 ities are in the Mississippi River diain- months, of adult males during spring age of Alexander County (Fig. 134). and summer molts, and of adult females Four additional collections of O. lan- during a summer molt. cifer recently have been made at Horse- The largest O. kentuckiensis from shoe Lake, where the species probab- ly is but difficult to collect. Illinois is a 37.5-mm CL form I male common collected on 25 April 1975; the largest Horseshoe Lake (Fig. 135) is an old. of the Missis- female is 36.2 mm CL, collected on 30 relatively deep oxbow September 1975. Both are from Big sippi River supporting stands of bald Creek. cypress and man> species of Coastal Plain animals that are rare in Illinois lancifer Orconectes (Hagen) (E\ers &: Page 1977). Few oxbows of the (Fig. 132) Mississippi River exist in southern Cambarus lancifer Hagen 1870 Illinois, and O. lancifer is unlikely to Cambariis faxonii Meek 1894 be found elsewhere in the state. Wolf Description. — Rostrum deeply ex- Lake in Lnion County appears to offer cavated; acumen longer than basal mar- suitable habitat for O. lancifer. but gin of rostrum, turning upward ante- several searches there ha\e failed to riorly; no median carina; margins con- document its presence. vex, diverging anteriorly, terminating In Louisiana. O. /fl;« //cr li\es in deep in spines. Carapace slightly flattened (more than 40 cm), clear, permanent August 1985 Crayfishes and Shrimps of Illinois ^32.-Orconecfes lancifer. A, dorsal view r J'9, of carapace; B. dorsal view of riaht chela- ---' -«- °' 9-opod of form ?'lTl'itT\Z':;,°'. I male; E. lateral v": o gonopod of form I male; F, lateral view of gonopod of form II male. 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 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. 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 Fig. 137—Total distribution of Orconectes attached. The eggs average 1.5 mm in placidus. 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 red fingers of ly brown; Fig. 138.— Distribution of Orconectes placi- chelae with red tips, subdistal black dus in Illinois; black dots = 1972-1982 col- bands. 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, Variation in O. placidus and its rela- populations herein assigned to O. tives is badly in need of study. placidus arc the most troublesome tax- Distribution and Habitat.— O. placi- onomically. The only large population dus occupies streams of the Cumber- is that in Big Creek in Hardin County; land, Tennessee, and lower Ohio rivers other populations are small, highly in Kentucky, Tennessee, and Illinois localized ones inhabiting rocky areas (Fig. 137). It lives in rocky riffles and along the banks of the Ohio and Missis- pools, using cavities and interstices as sippi rivers. Although all are assigned hiding places. to O. placidus, these populations show Recent Illinois records exist for Big variations usually indicative of specific Creek, Hardin County, the Ohio River, distinctiveness in Orconectes. The Massac and Pulaski counties, and the gonopod of the form I male lacks any Mississippi River, Randolph and Jack- hint of a shoulder on its dorsal surface son counties (Fig. 138). Most of the in Big Creek specimens but has a defi- large-river collections were made along nite shoulder (although not as pro- rocky banks or in rocky backwater areas. nounced and angular as in O. rusticus) In the Big Creek system, O. placidus is in Mississipi^i Ri\er specimens (no form found mainly in downstream gravel I males from the Ohio River are avail- and rubble riffles (Fig. 139), and O. able). Big Creek specimens always lack kentuckiensis is found mainly in up- a carina on the rostrum; those from the stream shallow rocky pools (Boyd &: Ohio and Mississippi rivers may or may Page 1978). As discussed below for O. not have one. In general, western j)op- rusticus, Hagen's (1870) and P'orbes' ulations differ the most from typical (1876) reference to "C. placidus" at O. placidus occupying the Cumberland Quincy is thought to refer to a waif of and Tennessee river systems in Ken- the Missouri po{)ulalions of O. ( f . rusti- tucky and Tennessee (Hobbs 1971b). 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 Number of Month Collections Fovm I Males Ovigerous Females Females with Young January 3 67 February 1 March 6 83 17 I April 2 50 May 6 June 19 July 3 August 1 September 7 71 October 8 88 November 4 75 December 1 100 The largest pieser\ed Illinois male is es ( = Form I) <: 1 r^MJu :;: a 45.6-mm form II; the largest female a is 41.7 mm CL. Both were collected in 5 10 Females May and, compared with the size-fre- quency distribution of October-col- " 5 o lected specimens (the largest sample), 1 ^ n n were probably 2 or 3 yearsold(Fig. 140). ^ . 10 20 30 40 The largest form I male (39.6 mm) has mm CL extremely large chelae (right chela = Fig. 140.—Size-frequency distribution of 4(>. 1 mm long). Among the Big Creek Orconectes placidus collected in October in specimens were 344 males and 387 fe- Big Creek, Hardin County. males (a ratio of 1:1.13). Among a combined total of 506 stripe- tail and spottail darters {Etheosloma rusticus" refers to the Big Creek popu- kennicotti and E. squainiceps) exam- lation of O. placidus. ined, the ingested remains of crayfishes Life History. — During the life- were found in the stomachs of 4 E. history study of O. kentuckiensis {Boyd squainiceps (Page 1974a, 1975). Con- & Page 1978), a large number (61) of sidering the relative abundances of cray- collections of O. placidus were made. fishes in Big Creek, these were almost The following observations are based certainly O. placidus or O. kentuckien- on those collections. sis (Boyd & Page 1978). Form I males were present in collec- tions from September through January and in March (only one collection was Orconectes propinquus (Girard) (Fig. 141) made in February) (Table 15). Brown (1955) found form I males in Big Creek Carnbarus pwpitiquus Girard 1852 in November. Orconectes iowaensis Fitzpatrick 1968 Ovigerous females (N = 'i) were found Description. — Rostrum exca\ atcd. only in March and April (Table 15); deflecting downward anteriorly; well- they were 15.7, 20.5, and 27.2 mm CL, developed median carina; long acumen; with 12, 88, and 134 eggs attached, margins thick, straight or slightly con- respectively. Eggs were spherical and tavc, barely coinerging anteriorly, ter- averaged 2.1 mm in diameter. No fe- minating in spines or lagc tubercles. males with young attached were found. 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- Fig. 142. —Total distribution of Orconectes pinquus ranges from southern Ontario propinquus. The northeastern limit of its range is unknown. 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 Fig. 143.— Distribution of Orconectes pro- the presence there of t)lher species of 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 landscape. It does best in clean, clear Orconectes. streams and presumably has suffered O. proptuquus may dig under stones population declines as stream siltation on the stream bed to escape desiccation has progressed during tfie last few dec- (Bovbjerg 1952; Brown 1955) or occa- ades. Among the widespread crayfishes sionally into stream banks (Hay 1896; of Illinois, it is probably the best in- Cralley 1932). Young often are found dicator of environmentally healthy in vegetation along the margins of a streams. stream. Altliough primarily an inhabi- All earlier records of O. propinquus tant of rocky streams, it can become in Illinois fall within, or almost within, established in lakes (Bovbjerg 1952). the present distribution. The record lor O. propinquus remains u'idespiead the Kaskaskia drainage, near Hum- and locally common, even abundant, boldt, Coles County (Rietz 1912), is one in some areas but probably was more county to the south of recent records. widespread and common prior lo the This record is signifitani in document- extensive modification of the Illinois 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. 146. —Presumed native distribution of (Fig. 145) Orconectes rusticus. Cambarus rusticus Girard 1852 Cambarus juvenilis Hagen 1870 O. rusticus by Bouchard (1977), O. Description. — Rostrum excavated; no juvenilis. median carina; long acumen; margins Distribution and Habitat. — O. rusti- thick, concave, terminating in spines. cus now occurs in New England. On- Carapace flattened dorsoventrally, with tario, Michigan, Ohio, Indiana, Ken- cervical spines; suborbital margin tucky, Tennessee, Illinois, Wisconsin, smooth. Areola moderate, narrowest Minnesota, Iowa, Missouri, and New part about 10-15 percent of length. Mexico. It has been widely introduced, Chelae large, heavily punctate; palm presumably through use as fishing bait, with two rows of tubercles on mesial and the boundaries of its native range margin. Form I gonopod with an an- (Fig. 146) now are difficult to determine. gular (90° to axis of gonopod) shoulder However, at least its occurrences in New on dorsal surface, terminating in two England (Crocker 1979), Ontario long elements: a sclerotized central (Crocker & Barr 1968). Wisconsin projection curved at its tip, and a slight- (Capelli & Munjal 1982). New Mexico ly shorter unsclerotized mesial process. (Bouchard 1977). and Illinois are cer- Dorsally green brown to rust red; often tainly the results of introductions. Once a large red spot surrounded by light introduced, O. ru5t?fU5 successfully dis- brown on side of carapace; fingers of places native species and expands its chelae with red tips, subdistal black range, apparently because it is a large bands. and aggressi\e species able to thrive Close relationships to O. rusticus are in impoundments and other disturbed not readily apparent. Species often habitats (Berrill 1978; Capelli 1982; confused with, and at times considered Capelli & Munjal 1982: Capelli & related to, O. rusticus include O. for- Magnuson 1983). Its continued use as ceps, O. placidus, O. putnami, O. a li\e bait species is therefore unwise. spinosus, and until synonymized with Taxonomic confusion about O. rus- August 1985 Crayfishes and Shrimps of Illinois 413 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) Fig. 147. — Distribution of Orconectes rusti- cus in Illinois. Open circle = Adams County and probably is supplanting native locality given by Hagen (1870) for "Cambarus species, as it is in southern Ontario placidus," black dots = 1972-1984 collections. (Berrill 1978) and Wisconsin (Capelli & Munjal 1982). Hybridization between O. rusticus and O. propinquus and ticus and its relatives has reigned since between O. rusticus and O. limosus their descriptions. "Cambarus wiscon- has been recorded in Wisconsin and sinensis Bundy," found at Normal Massachusetts (Capelli & Capelli 1980; (Forbes 1876), is a synonym of O. virilis Smith 1981) and appears to represent (Hobbs 1972a) and is not a reference to an intermediate step in the displace- O. rusticus, as thought by Rietz (1912) ment by O. rusticus of these native and Page (1974b). Brown's (1955) species. In Illinois, it appears to be records for "O. rusticus" all were for hybridizing with O. virilis in Wolf Big Creek in Hardin County and are Lake. O rusticus lives in rocky riffles, treated herein as O. placidus. pools, and along wave-swept lake "Cambarus placidus Hagen" was margins. recorded from Quincy by Hagen (1870) Life History. — In fish ponds in Ohio, and Forbes (1876) and considered by Langlois (1935) found mating to occur Rietz (1912) to represent a record of in September and October; following O. rusticus. Quincy is directly across copulation, the females burrow into the Mississippi River from the Fabius banks. Oviposition occurs in a few River system of Missouri which sup- females in October, but mostly in April ports a population of O. rusticus or a and May, when females are 1 year old. closely related undescribed species. The Eggs hatch in about 20 days and remain Quincy record probably represents a attached to the female for al)out 5 days. waif from the Fabius River population 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. subparallel, slightly converging me- Prins (1968) studied O. rusticus in a dially, terminating in spines. Carapace stream in Kentucky and found that flattened dorsoventrally, with cervical mating peaks in September and Octo- spines; suborbital margin weakly an- ber, females oviposit from February gular. Areola moderately wide, narrow- through June at about 22-24 months, est part about 15-25 percent of length. eggs hatch in May after being carried Chela large, heavily punctate; palm by the female for 4-6 weeks, young with two rows of tubercles on mesial remain attached to the mother for 3 margin. Form I gonopod terminates weeks {through April and May), sexual in two short, straight elements: a sclero- maturity is reached at about 15 months tized central projection and an unsclero- (about 18 mm CL), the sex ratio among tized mesial process with spur about juveniles is 1:1, and the species reaches middle of caudal surface; shoulder on a maximum longevity of about 3 years cephalic surface at base of central pro- and a maximum CL of 45 mm. Counts of jection. Olive green to light brown and ovarian eggs in 43 females ranged from usually heavily speckled with dark 54 to 357. The smallest female with eggs brown dorsally; fingers of chelae with attached (55 eggs) was 17 mm CL; _the red tips, subdistal black bands. largest, with 231 eggs, was 38 mm CL. Distribution and Habitat. — O. stan- The relationship between the number nardi is a member of the O. propincjuus of eggs (N) and carapace length (L) species group that is endemic, and 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, and the largest female is 41.5 mm CL. Orconectes stannardi Page (Fig. 148) Fig. 149— Distribution of Orconectes stan- Orconectes stannardi Page 1985 nardi. Open circles = localities given by Rietz excavated; Description. — R(jstrum (1912) and Brown (1955) for "O. propinquus" well-developed median carina; long and "O. indianensis." black dots = 1972-1984 acumen; margins moderately thick. 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- 1 August 1985 Crayfishes and Shrimps of Illinois 417 corded localities for O. propinquus and tains two ovigcrous females. All 20 O. indianensis in the Little Wabash males collected in November, 5 of 1 River system that almost certainly were collected in August, and 1 of 25 collected based on misidentified O. slannardi in May are form I. The two ovigerous (Page 1985). Data collected during the females collected in May 1984 were 20.7 present survey suggest that O. pro- mm and 28.9 mm CL and carried 124 pinquus, O. indianensis, and O. stan- and 184 eggs, respectively. Eggs average nardi occur allopatrically and that 1.9 mm in diameter. The largest speci- neither O. propinquus nor O. indianen- men is a 38.3-mm CL female collected sis occurs in the Little Wabash River on 22 August 1984; the largest male, system. Unfortunately, the collections from the same collection, is a 30.5-mm made by Rietz and Brown cannot be form L located and re-examined. Although all collections of O. stan- nardi that have been made since 1972 are Orconectes virilis (Hagen) from the Little Wabash River proper, (Fig. 151) some of the collections of Rietz (1912) Cambarus virilis Hagen 1870 and Brown (1955) were from tributaries. Carnbarus debilis Bundy 1876 Collections of Rietz (1912) were made in Cambarus wisconsinensis Bundy 1876 Big Muddy Creek between Richland Cambarus couesi Streets 1877 and Clay counties and in Skillet Fork Description. — Rostrum excavated; at Wayne City, Wayne County; the moderately long acumen; no median former was identified by Rietz as O. carina; margins thick, straight, barely propinquus and the latter as O. in- converging anteriorly, with terminal dianensis. Collections cited by Brown spines. Carapace flattened dorsoven- (all as O. propinquus but which must trally, with cervical spines; suborbital have been of O. slannardi) were made margin bluntly angular. Areola narrow, in Dismal Creek, 2\ mi N lola. Clay constricted anteriorly, narrowest part County; Salt Creek, 1 mi SE Effingham, about 6-7 percent of length. Chelae Effingham County; Blue Point Creek, large, heavily punctate; palm with two 2'2 mi S Shumway, Effingham County; rows of tubercles on dorsomesial mar- Lost Fork, I'j mi E Omega, Marion gin. Form I gonopod terminates in two County; branch of Little Wabash River, long, caudodistally directed elements: 1'4 mi SW Trowbridge, Shelby County; a sclerotized central projection and and Little Wabash River, 4 mi NE a shorter unsclerotized mesial process. Shumway, Effingham County. Dorsally Hght brown to dark green- Most individuals were found in shal- brown; fingers of chelae without bright low riffles composed of large flat stones red tips and subdistal black bands. (Fig. 150). A few were found in deeper O. virilis occasionally occurs as a water but always in association with blue color morph in Illinois and else- stones or accumulations of sticks and where. Sugar Creek, a tributary of the other debris. The preference of O. Sangamon River in McLean County, slannardi for rocky riffles is typical contains a large number of blue O. of the Propinquus group of Orconecles, virilis. The upper reaches of Sugar the members of which typically hide Creek are highly modified and receive under stones and among debris. O. a large amount of pollution from slannardi is locally common in the Bloomingion-Normal. and ihe large Little Wabash River. number of blue crayfish may somehow Life History. — All seven collections be related to the degradation of the of O. slannardi presently available stream. Anderson (1975) descrilxxl a (from November 1973, Nf)vember 1983, 4 1 -mm C;i, female O. virilis from the May 1984, and August 1984) contain Kisliwaukie Ri\er at De Kalb. Illinois, form I males, and one (May 1984) con- 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 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. 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, Fig. 153.— Distribution of Orconectes virilis exists in huge populations. It occurs 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 Cralley (1932) found numerous bur- fantail darter, Etheostoma flabellare; rows belonging to O. virilis opening and the green water snake, Nerodia just above the water line along the banks cyclopion (Webb, Penny, & Marlin of the Kaskaskia Ri\er in Champaign 1975; Smith 1961, 1979). County. Brown (1955) found a few A recent record for O. viriUs in an burrows of O. virilis in Illinois and impoundment in the Saline River sys- described them as being simple, with an tem (Berkel Lake, Saline County, 9 enlarged pocket at the lower end and October 1977) is almost certainly the only one surface opening. Phillips result of a recent introduction. The ( 1980) found a burrow in Iowa about 30 record is based on two individuals, one cm deep with two terminal chambers of which is small (16.3 mm CL), sug- and a chimney about 5 cm tall. Hazlett gesting that the species is reproducing et al. (1974) found O. I'irilis burrowing in the lake. extensively along a stream and pond in In Illinois, O. virilis occupies rocky Michigan; burrows frequently had two streams (Fig. 154) and lake margins. It entrances, often one above the water usually occurs in moderate current level. Rarely during the present survey but is less common in swift riffles was O. I'irilis found in burrows, and it (where, throughout much of Illinois, appears to be much more common for it is replaced by O. propinquus) and O. virilis to hide under rocks, even in in sluggish pools (where O. inunimis winter, than to burrow; however, as predominates). The largest populations discussed below, o\ igerous females may are found in areas where large flat stones habiiualh burrow. provide a preponderance of hiding Life History. —The wide-ranging and places. In lakes, O. virilis may occur often abundant O. i'irilis has been the as deep as 9 m (Crocker &: Barr 1968). 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). lected during the present sur\ey was Momot & Gowlng (1977) found much 1,243 males: 1,1 82 females. lower egg counts among high-density populations of O. virilis in Michigan Genus Fallicambarus Hobbs lakes. The number of attached eggs FaUicambarus Hobbs 1969 on 402 females (22-35 mm CL) aver- Gonopod with two large terminal 91.9. were aged only These females elements curved at about 90° angle to Illinois. smaller than those found in shaft; central projection large and Also, fecundity in these lakes was bladelike. Ischium of third pereiopod thought to be related to density- de- of male with hook. Opposable margin processes, i.e., as the density pendent of dactyl of chela with deep conca\ ity. the of eggs carried increased number Only F. fodiens of the 14 recognized by females decreased (Momot Sc Gowing species of Fallicambarus (Hobbs 1981) 1977). occurs in Illinois. F. fodieiis is assigned Elsewhere, form I males have been to the subgenus Creaserinus and is most found in April and June-October in closely related to F. hedgepethi (see New England (Crocker 1979), May and comment below) and F. uhleri (Hobbs in (Crocker August New York 1957), 1973). June-August in Ontario (Crocker & Barr 1968), and July-April in Iowa Fallicambarus fodiens (Cottle) (Caldwell & Bovbjerg 1969). Form I (Fig. 155) males may be as small as 25 mm CL (Berrill 1978; Weagle 8c Ozburn 1972). Astacus fodiens Cottle 1863 Females carrying eggs have been Cambarus argil licola Faxon 1884 found in April and May in Iowa (Cald- Description. — Rostrum broad, ex- well Sc Bovbjerg 1969; Phillips 1980), cavated, with short acumen, lacking April and May in Michigan (Pearse marginal spines and tubercles, lacking 1910; Momot & Gowing 1977) and New median carina. Carapace compressed, Hampshire (Crocker (1979), May in lacking cervical spines; suborbital Rhode Island (Crocker 1979), and May- margin smooth (not angular). ,\reola July in Ontario (Crocker Sc Barr 1968). obliterated. Chela large, punctate; Females with young attached have been palm with one to three rows of tubercles collected in May in Iowa (Phillips 1980) on mesial margin; dactyl with deep and in June in Maine (Crocker 1979). concavity on basal half of opposable Some Illinois specimens appear to be margin. Form I gonopod terminates among the largest recorded, although in two elements, both cur\ ed at an angle Creaser (1932) noted that in Wisconsin of about 90° to the principal axis of the species "frequently attains a size the gonopod: a distally rounded sclero- of over eight inches" (about 100 mm tized central projection and a short CL). The largest Illinois specimens proximally thick mesial process with a are from reservoirs: a 59.5-mm CL fe- pegiike tip. Dorsal cx:)loi variable, usu- male from Dawson Lake, McLean ally olive brown with dark brown County, collected on 25 September blotches, but often a uniform light 1972, and a 59.2-mm CL form I male brown to rust red. from Baldwin Lake, Randolph County Distribution and Habitat. F. fo- collected on 2 November 1972. O. mrilis difns is the most wide-ranging species is an agressive crayfish and, with its in its genus, extending, in the Great large size giving it an advantage, ac- Lakes (except Lake Superior) and Mis- tively excludes the often syniopic O. sissippi Ri\er drainages, from souiiiern iniDnniis from desirable hiding places Ontario to wesieni Kentuckx (Fig. 156). (Bovbjerg 1970). Intiaspecific domi- South of Kentucky it is replaced by the nance was examined by Bovbjerg (1953). clo.sely related, and probablv conspe- The sex ratio among specimens col- 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. Fig. 157.— Distribution of Fallicambarus personal communication), F. Iiedge- fodiens in Illinois. Large circles = 1890-1901 pethi, which ranges to the Gulf Coast collections; small circles = collections made in and into Gulf drainages from Georgia the 1950's; black dots = 1972-1982 collections. to Texas. In Illinois it is known from the Ohio (including the Wabash), Big Muddy, -Ottawa, LaSalle Co., 111.. Sept. 1901," Sangamon, Illinois, and Lake Michigan on Bovbjerg's (1952) study of F. fodiens drainages (Fig. 157). It appears to be and O. propinquus in the Lake Michi- absent from western and extreme north- gan drainage of southern Cook County, ern Illinois and is rare in central Il- and Bovbjerg's (1952) mention of the linois. Although difficult to collect presence of F. fodiens in Dead River at because of its burrowing habits, present Zion, Lake County, also a tributary of F. records suggest that it remains common Lake Michigan. Brown (1955) found in southern Illinois in the Ohio, Big fodiens in a woodland pond (Sangamon Muddy, Saline, and Little Wabash River system) 2.5 km SW Langley\ille, systems. Christian County. During the present survey. F. fodieiis In Illinois, F. fodiens lives on wooded was collected in central Illinois only floodplains (Fig. 158) or wheie wooded on the Sangamon River floodplain at floodplains formerly occurred. It lives Allerton Park in Piatt County; however, in deep burrows and is in surface waters historical records suggest that the (usually floodplain ponds) only dining species was formerly more common in floods: 27 of .'51 collections of F. fodiens central and northern Illinois. The made in Illinois since 1972 were made northernmost records shown in Fig. from February through May, the period 157 are based on a collection at the Il- of late winter-spring flooding, although linois Natural Historv Survey labelled 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 Form I) 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 Fig. 159.—Size-frequency distribution of Fallicambarus fodiens collected in April in & Barr 1968). 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 in Ontario with eggs in March and typically have one to three entrances, April and young in May (Crocker & Barr often capped by chimneys, leading to 1968), and in Illinois with eggs in Feb- an oval resting cavity (Faxon 1885) and ruary and March and young in Feb- leading down from the cavity 30-60 cm ruary, March, and April (Table 18). is a narrow escape tunnel (Crocker & Brown (1955) dug a female with at- Barr 1968). Soils having a high clay tached young from a burrow in south- content are probably necessary for the ern Illinois in October. deep burrows constructed by F. fodiens In the present survey, three females (Crocker & Barr 1968). with eggs and 24 females with attached Life History. — In a study in southern young were collected. The females with Cook County, Illinois, Bovbjerg (1952) eggs were 22, 28, and 35 mm CL, with found juveniles and gravid females in 48, 68, and 162 eggs, respectively. Eggs woodland ponds as early as March. By on the two smaller females, one col- June the adults had disappeared, pre- lected each in February and March, sumably into burrows, and only young averaged 2.2 mm in diameter; those on remained in the ponds. By 2 weeks prior the largest female, collected in March, to the drying of the pond, all crayfish averaged 2.5 mm. Seven females col- had disappeared from the pond, either lected in February and March (30-44 by burrowing or emigrating. Individ- mm CL) had 96-201 (mean, 132.1) uals were found (one at a depth of 1.2 young attached. m) throughout the summer by digging The size-frequency distribution of in the dry pond bed down to the water southern Illinois, April-collected F. table. The refilling of the pond in the fodiens suggests that the species reaches fall was followed by a reinvasion of sexual maturity and dies in its second F. fodiens, which was active in the pond year. Only males over 40 mm CL (second in November under a 5-cm ice cover. year) among the indi\iduals collected F. fodiens can survive out of water were form I, and all females over 30 mm (at 20.5° C and 66 percent relative hu- CL (second year) were carrying young. midity) for about 12 hours (Bovbjerg All smaller males and females were 1952), and overland migrations of F. immature (Fig. 159). The sex ratio fodiens on warm (>5°C) humid among the combined April samples was nights in March-May were docu- 44 males: 42 females. mented by Cummins (1921). F. fo- diens is omnivorous, and in northern Genus Cambarus Erichson Illinois specimens, stomach contents included plant fragments (especially Carnbarus Erichson 1846 grass seeds), insect fragments, crayfish Bartonius Ortmann 1905 appendages, an isopod, and a sala- Gonopod with two large terminal mander larva (Bovbjerg 1952). elements curved at about 90° angle to Sexually mature (form I) males occur shaft; central projection large and in March and April in Michigan (Cum- bladelike. Ischium of third pereiopodof mins 1921) and in April (Table 18), male with hook. Opposable margin of August, and October (Brown 1955) dactyl of chela without deep concavity. in Illinois. The two form I males col- Cambarus contains about 77 species lected during the present survey had (Hobbs 1981), of which four occur in carapace lengths of 42 and 43 mm. Illinois. Two of the Illinois species In Michigan, females have been (lenebrosus and rusticiformis) belong found with eggs in March and April to the subgenus Erebicambarus, one and with young in April and May (dio^enes) to Lacunicambarus. and one (Pearse 1910; Cummins 1921: Creaser (robuslus) to Punclicambarus. The 1931), in Indiana with eggs and young wide-ranging C. diogenes appears to be in April (Hay 1896: Williamson 1907). a complex of species (Hobbs 1974b), August 1985 Crayfishes and Shrimps of Illinois 427 Fig. 160— Cambarus tenebrosus. 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 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 Fig. 161.—Total distribution of Cambarus terminates in two elements, both curved tenebrosus. 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 Fig. 162, —Distribution of Cambarus tene- brevis (Rietz 1912), C. bartoni laei'is = brosus in I linois; black dots 1972-1982 col- (Brown 1955), C. laevis (Hobbs 1968, lections. August 1985 Crayfishes and Shrimps of Illinois 429 1969, 1972a; Page 1974b: Peck & Lewis the Shawnee Hills of southern Illinois, 1977), C. ornalus (Peck & Lewis 1977), into the Cumberland River drainage of and C. teuebrosus (Peck & Lewis 1977). western Kentucky, and into the general It now appears that the correct name for range of C. teuebrosus with no notice- the southern Illinois populations is C. able mor|)hological discontinuities. teuebrosus. Populations extend across 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; annulus C, 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 Fig. 165.—Total distribution of Cambarus 1983) suggests that "when adequately rusticiformis. 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 Fig 166— Distribution of Cambarus rustici- = 1975 collection. in Hardin County fed by numerous formis in Illinois; black dot — 432 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 springs (Fig. 163). Elsewhere in Illinois CL was found to be about 65 mm and it has been found only in Willow Creek maximum longevity about 36-38 (Vermilion River system) in Vermilion months. The sex ratio among 3,235 County, Illinois (Fig. 162), where it specimens was 1 male: 1.03 females. presumably is distributionally contig- No form I males or females carrying uous with the Indiana population. eggs or young were present among the The Illinois and total ranges of C. 40 collections of Illinois C. tenebrosus lenebrosus are quite similar to those made during the present survey (all of the zigzag salamander (Plethodon months except September). Brown dorsalis) although the salamander has (1955) reported collecting form I males dispersed into Clear Creek, a tributary in August and October and a female of the Mississippi River, in Alexander, with attached young in October. All Union, and Jackson counties (Smith of Brown's collections were made in 1961). Hardin and Pope counties, Illinois. C. tenebrosus, first recorded in Illi- A 46.6-inm CL female from Big Creek nois (as C. bartoni brevis) by Rietz is the largest Illinois specimen. (1912), probably has essentially the same distribution now as it had prior to the Cambarus rusticiformis Rhoades arrival of man. However, the water table (Fig. 164) of southern Illinois has been lowered within historic times, and some springs Cambarus rusticiformis Rhoades 1944 may have disappeared or at least are Description. — Rostrum narrows smaller than before, reducing the habi- anteriorly; margins elevated, concave, tat available for C. tenebrosus. Such and tipped with large knobs; acumen reductions in habitat have caused some moderate, ele\ated anteriorly; median spring-inhabiting fishes (Phoxinus carina lacking. Carapace dorso\entrally erythrogaster, Rhinichlhys atratulus) flattened, with large cervical spine, to disappear from southern Illinois small suborbital angle. Areola mod- (Smith 1979). erately wide, narrowest part 16-18 per- Life History.— Prins (1968) studied C. cent of length. Chela large. hea\ily lenebrosus in a stream system in Ken- punctate; palm with single row of tucky and found it to be most common flattened tubercles along mesial surface. in constant-temperature headwaters. Form I gonopod terminates in two ele- Most adults were found in burrows or ments, both directed caudalK at an under large rocks, but young and sub- angle greater than 90° to the principal adults were found in open-stream axis of the gonopod: a distally rounded, environments. The diet consisted pri- sclerotized central projection and a marily of vascular plants, especially large distally tapering mesial process. fallen leaves from the riparian habitat, Distribution and Habitat. C. rusti- and filamentous algae. Other food items ciformis occupies the Cumberland included detritus, aquatic insects, and River drainage and the Barren Ri\er crustaceans. Hatching occurs in July system (Green Ri\er drainage) of Ken- and August after eggs have been carried lucky and Tennessee (Bouchard 1976) by the female for at least 25-30 days. and, judging from one Illinois collec- Growth continues throughout the year, tion, the lower Ohio River (Fig. 165). including winter, and sexual maturity Two juvenile males (11.7 and 14.6 mm is reached at about 20-22 months and CL) were collected in a rockv backwater 10 mm CL. Mating occurs in winter of the Ohio Ri\er near Lock and Dam and spring. Mature Irmales oviposit 53, 4 km NE Olmsted, Pulaski County, in June and July at about 22-23 months Illinois, (Fig. 166) on 16 July 1975. and probably again at 34-35 months. Several later attempts to find C. rusti- Egg counts on seven females (44-52 mm ( iformis at the same and other sites on (;L) ranged from 186 to 266. Maximum the lower Ohio Ri\er were imsuccessful. August 1985 Crayfishes and Shrimps of Illinois 433 diogenes. A, dorsal view r ^nL^V^^^f^!"^^/;"^ of carapace: B, dorsal view of right chela- ^'"^ aonopod of ",°' °* form I male: E, lateral view of gonopod oi formPnr^T^^LTI male: F. lateral'f "^^n" view of gonopod of form II male 434 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 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. 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 Fig. 169.— Distribution of Cambarus diog- Commission on Zooligical enes in Illinois; blacl< dots = 1972-1982 col- Nomenclature, Opinion 522, 1958.) 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 mont and Coastal Plain from New green with red outHning abdominal Jersey and Pennsylvania south to cen- segments, tips of chelae, and margins of tral Georgia, in Gulf drainages from rostrum. Georgia to eastern Texas, and north Marlow (1960) examined geographic in the Mississippi and Great Lakes variation in C. diogenes and concluded basins to western New York and Penn- that only two subspecies could be rec- sylvania, Michigan, Wisconsin, Min- ognized: C. d. ludovicianus in the Lake nesota, and North Dakota. The western- Ponchartrain drainage and southern most populations occur in the Platte Louisiana and C. d. diogenes through- Ri\er drainage of Colorado and Wy- out the rest of the range of the species. oming (Fig. 168). However, according to Hobbs (1969, C. diogenes occurs statewide in Il- 1974b), C. diogenes is a complex of linois (Fig. 169) and is generally com- forms, and C. d. ludovicianus probably mon. It is abundant in the Shawnee is more widespread than Marlow Hills and on the Coastal Plain of south- thought. Illinois specimens show tre- ern Illinois, less common in northern mendous variation in the shape of the and western Illinois. Its burrowing carapace and rostrum and in coloration. habits preclude easy collecting, and If several taxa presently are included in consequently, the number of (olle( lions "C. diogenes." more than one may be (Fig. 169) does not adetjuateh reflect found in Illinois. The only other de- its abundance in Illinois. scribed species in the subgenus Lacuni- C. diogenes first was recorded from cambarus are C. acanthura, restricted Illinois by Hagen (1870), who described to the Tennessee and Alabama drain- the spec imens (ollected at Lawn Ritlge ages of Tennessee, Alabama, and (ieoi- (Marshall County), Belleville, and gia (Hobbs 1981) and C miltus from Fvanston as a new species, Cambarus Baldwin Clounty, Alabama (Filzpatritk obesus. Forbes' (1876) only comments 1978b). on "('.. obesus" were: "Very tommon. Dislribulion and Habitat. C. diog- The largest in the state." enes ranges along the Atlantic Pied- 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 of torm 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 ruary-April) and in fall (September- July (except in May), and in October November). and November (Table 19); Brown (1955) Ovigerous females have been found found form I males in Illinois from in January (Bundy 1877), April (Hay March through September, except in 1896; Evermann &: Clark 1920: Creaser July. The primary peak of occurrence 1932), and May in Indiana (Williamson is in February and March, and a second- 1907); in April and May in Pennsyl- ary peak occurs in November (Table 19). vania (Ortmann 1906) and Iowa (Phil- Elsewhere, matings have been observed lips 1980); in April (Williams & Leonard in March, April, May (Hargitt 1890; 1952) and May in Kansas (Harris 1900); Hay 1896; Evermann Sc Clark 1920), and from late March to May in Wisconsin October (Williams & Leonard 1952). In (Graenicher 1913); and in September in ). Louisiana, form I males were found in Georgia (Hobbs 1981 Marlow ( 1960) March, April, and October (Penn &: examined o\igerous females from Marlow 1959) and in Georgia in Jan- March (1 female), April (19), May (17), uary, April, and October (Hobbs 1981). and October (I). Females carry young Ortmann (1906) recorded form I males in May and June in Pennsylvania in Pennsylvania from March through (Ortmann 1906) and in Indiana (Wil- November. Among the C. diogenes liamson 1907; Evermann &: Clark 1920), examined by Marlow (1960), form I and in June in Michigan (Creaser 1932). males were present in every month ex- In Illinois, females were found to carry cept January, but were most prevalent eggs from January through Ma\, except 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- Fig. 173, — Distribution of Cambarus robus- tus in Illinois. The two records (Adams and Fig. 172.— Total distribution of Cambarus Macon counties) are from Faxon (1885) and robustus. Rietz (1912). August 1985 Crayfishes and Shrimps of Illinois 439 in February, and young in May and North Carolina and eastern Kentucky June (Table 19). and west to Michigan and central In- Because of the difficulty in obtaining diana (Fig. 172). It occupies rocky areas adult C. diogenes, few females with of fast-flowing streams, and sometimes eggs or young attached were collected is found in rocky lakes and ponds in the present survey. Five females (Crocker &: Barr 1968; Berrill 1978). (43.7-56.4 mm CL) collected in April The only Illinois records (Fig. 173) and May were carrying 183-518 (mean, are from the late nineteenth and early 295.2) eggs. The highest number of eggs twentieth centuries. Faxon (1885) re- was on the smallest female. Eggs varied ported C. robustus from Decatur in diameter from 2.1 to 3.0 mm, and (Macon County, Sangamon River sys- unlike other crayfishes examined dur- tem), and Rietz (1912) reported four ing the present survey, in C. diogenes specimens from Quincy (Adams egg diameter has a significant corre- County, Mississippi River drainage). lation (r = 0.97) with length of the fe- Although the Quincy specimens cannot male: egg diameter = -1.280 + 0.077 be located, the Decatur specimens are CL. in the U.S. National Museum (USNM A 48.5-mm CL female collected in, 63291) and unquestionably document Johnson County on 19 May 1973 had the former presence of the species in 606 early-instar young attached; a 47.2- the state. C. robustus is rare even as far mm CL female collected in Christian west as Indiana (Eberly 1955), and the County on 8 June 1979 had 45 late-instar Illinois collections must have repre- young attached. Two females collected sented relict populations left over from in Indiana in 1906 were carrying 87 a more widespread distribution in a and 215 young (Williamson 1907). cooler postglacial climate. Many streams today around Quincy are Cambarus robustus Girard spring-fed, and considerable effort was (Fig. 171) expended in unsuccessful attempts to Cambarus robustus Girard 1852 locate surviving populations of C. Description. — Rostrum moderately robustus in what appear to be suitable broad, shallowly excavated, lacking habitats. The Sangamon River, flowing median carina; margins converge an- through Decatur, was badly polluted teriorly, lack terminal tubercles or in the early part of this century (Jewell spines; acumen moderately large. Cara- 1920), is now impounded by Lake De- pace dorsoventrally flattened, with catur, and appears unlikely to support large cervical spines; suborbital margin a population of C. robustus. angular. Areola moderately wide, nar- Life History.— Crocker (1957) re- rowest part about 12-17 percent of ported form I males in New York col- length. Chela large, heavily punctate; lections made from April through Octo- palm with two rows of tubercles on ber and observed copulation in October mesial surface; base of fixed dactyl and May (the latter in a lamprey trap). with deep dorsal and ventral imjjres- Form I males have been collected in in and sions. Form I gonopod terminates in Pennsylvania May July-No- two elements: a distally rounded, sclero- vember; the smallest was about 63 mm tized central projection curved at an total length (about 32 mm CL) (Ort- angle of more than 90° to the principal mann 1906). In southern Ontario, form axis of the gonopod, and a large gently I males were 25-55 mm CL (Berrill tapering mesial process curved at about 1978). Ovigerous females have been 90° to principal axis. Dorsally green found in July in New York (Crocker brown. 1957) and Pennsylvania (Ortmann Distribution and Habitat. — C. ro- 1906); the Pennsylvania female, 84 mm bustus ranges from Connecticut and total length (about 42 mm CL), carried southern Ontario south to northwestern 228 eggs. Females with young attached 440 Illinois Natural History Survey Bulletin Vol. 33. Art. 4 have been observed in April and August occurs only in a small area of Indiana. in New York (Crocker 1957). Young feed Orconectes kentitckiensis occurs only on immature insects; adults feed mainly in Big, Hosick. and Peters creeks in on aquatic plants (Greaser 1934). Hardin Comity, and although the dis- tribution of this species in Illinois CONSERVATION probably has changed little historically, its persistence is threatened by modifica- Unfortunately, it appears that Macro- tions of the riparian habitats in Hardin brachium ohione and Cambarus ro- County. Trees have been remo\ed from biistus already have disappeared from many miles of stream bank, resulting Illinois. C. robustus may have dis- in less shading of the stream and higher appeared from Illinois naturally rather water temperatures. Water has been than because of man-induced changes diverted in ever-increasing amounts in the environment. Our earliest records from its natural drainage for use in indicate that it was relict in Illinois, agriculture. The lowered water table presumably having been reduced from a is reducing the size of Big Creek. Else- more widespread distribution to small where, O. keutuckiensis occurs only in areas near Quincy and Decatur. Else- a small area of northwestern Kentucky. where it inhabits cool to cold streams Orconectes placidus is common and apparently was reduced in warm throughout much of Kentucky and postglacial Illinois (probably during Tennessee, but in Illinois it maintains the Xerothermic period) to small popu- a sizeable population only in Big Creek lations that subsequently were imable in Hardin County and is threatened by to perpetuate themselves. A7. ohione the same stream modifications threat- probably disappeared because of the ening O. kenluckiensis. extensive modification and degradation Orconectes lancijer is restricted in of Illinois' large rivers. Especially Illinois to Horseshoe Lake in .\lexander harmful loM. ohione would have been County. Although under the supervi- the loss of aquatic vegetation associated sion of the Illinois Department of Con- with increased siltation, channeliza- ser\ation, the lake has been severely tion, and chemical pollution. modified in recent decades, and much Unless protective measures are taken, of it now is surrounded by agricultural several more species of native decapods fields ami grasses. If the unusual biota of are likely to disappear from Illinois. Horseshoe Lake is to persist, subsequent The following four species have re- modifications of the lake shoidd be stricted ranges in the state and are allowed only after carefid consideration threatened with extirpation. They of their iin ironmental inipaci. Else- should receive as much protection as where O. lancifrr occurs imcoinmonly possible by our adding them to the ihoughout the former Mississippi Em- Illinois List of Endangered and Threat- bayment. ened Species and protecting their habitats. The distribution of Orconectes in- ACKNOWLEDGMENTS dianensis has been reduced dramatically I am extremely indebted to Mr. Craig in Illinois as a result of severe pollution W. Ronto for the execution of most of and other major modifications of the the figures appearing in this pidilica- Saline River system. Remaining popu- tioii and lo Dr. H. H. Hobbs, jr.. for lations are restricted to the South Fork laxononiic comisel, distributional in- of the Saline River in Gallalin, John- formation, and a thorough review of son, Pope, Saline, and Williamson the manuscript. (ounties, Robinette (Eagle) Creek in I am indebted also to B. M. Burr and (.allaiin Comity, and Honey Creek in R. W. Bouchard for taxonomic (inmsel ilavdin C^ounty. Elsewhere the species during the study, to J. .\. Beatty, P. W. August 1985 Crayfishes and Shrimps of Illinois 441 Smith, G. C. Sanderson, and C. Burnett Berrill. M. 1978. Distribution and ecology of for helpful reviews of the manuscript, crayfish in the Kauartha region of southern Ontario. Canadian Journal of Zoology 56:166- to Bernice P. Sweeney for typing the 177. manuscript, and to the following col- Black, B. 1966. Cyclic male reproductive activ- leagues and friends for assistance with J. ities in the dwarf crawfishes Cambarellus shu- field work and/or providing specimens: feldti (Faxon) and Cambarellus puer Hobbs. J. Allen, R. V. Anderson, N. A. Ashford, American Microscopical .Society Transactions 85:214-232. S. S. Birky, L. Boscarine, J. K. Bouse- man, J. A. Boyd, R. A. Brandon, A. R. . 1972. Life history irotes on the crawfish Brigham, W. U. Brigham, B. Britton, Orconecles lancifer. Louisiana Academy of Sciences Proceedings 35: 7-9. J. Brower, K. Brummett, J. Buerger, B. M. Burr, P. A. Burr, M. K. Butcher, Bouchard, R. W. 1976. Geography and ecology R. W. Clegern, L. E. Cordes, A. 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A general revision of the Decapoda Natantia) Schwartz. 1960. Paiaemonidae (Crustacea Meredith, W. G., and F. J. of the Americas. II. The subfamily Patae- Maryland crayfishes. Maryland Department Occa- Series moninae. Allan Hancock Foundation of Research and Education. Educational sional Paper 12. 46. 1967. Population dvnamics and HoRBERG, L. 1945. A major buried valley in east MoMOT, W. T. of the ciavfish. OKonecles iinhs. central Illinois and its regional relationships. productivity American Midland Natiiralisi Illinois C;eological Survey Report 106: 349-359. in a marl lake. 78:55-81. production and produc- . .\nnual HiiBSCHMAN. J. H., and J. A. Rose. 1969. Palar- 1978. Orconecles monetes kadiakni.sis Rathbun: Post embrvonic tion biomass ratios of the crayfish. lakes, .\merican growth in the laboratory (Decapoda, Palae- I'lrilis. in two norihcrn Ontario monidae). Crustaceana 16:81-87. Fisheries Society Transactions 107:776-784. August 1985 Crayfishes and Shrimps of Illinois 445 , and H. GowiNC. 1977. Production and . 1952. The genus Orconectes in Louisiana population dynamics of the crayfish Orconectes (Decapoda, Astacidae). American Midland virilis in three lakes. Michigan Fisheries Re- Naturalist 47: 743-748. search Board of Canada Journal 34: 2041-2055. . 1956. The genus Procambarus in Louisi- Newcombe, C. L. 1929. The crayfishes of West ana (Decapoda, Astacidae). American Midland Virginia. Ohio Journal of Science 29:267-288. Naturalist 56: 406-422. Nielsen, L. A., and B. Reynolds. 1975. Fresh- J. . 1959. An illustrated kev to the crawfishes water shrimp natural for fishes. food pond Farm of Louisiana with a summary of their distribu- Harvest Pond 9(2): 8,9.24. tion within the state (Decapoda, Astacidae). 3-20. , and . 1977. Population character- Tulane Studies in Zoology 7: istics of a freshwater shrimp. Palaemoneles and J. F. Fitzpatrick, Jr. 1962. Inter- kadiakensis Rathbun. Missouri Academy of specific competition between crawfishes. Amer- Science Transactions 10 & 11:44-57. ican Zoologist 2: 436. Ortmann, a. E. 1906. The crawfishes of the State , and . 1963. Interspecific competi- of Pennsylvania. Carnegie Museum Memoirs tion between two syinpatric species of dwarf 343-523. 2: crawfishes. Ecology 44: 793-797. . 1931. Crawfishes of the southern Ap- -, and H. H. Hobbs, Jr. 1958. A contribu- palachians and the Cumberland Plateau. Car- tion toward a knowledge of the crawfishes of negie Museum Annals 20:61-160. Texas (Decapoda, Astacidae). Texas Journal Pace, L. M. 1974a. The life history of the spottail of Science 10:452-483. darter, Elheostorna squamueps, in Big Creek, , and G. Marlow. 1959. The genus Cam- Illinois, and Ferguson Creek, Kentucky. Illinois barus in Louisiana. American Midland Nat- Natural History Survey Biological Notes 89. uralist 61:191-203. . 1974b. Aquatic Malacostraca recorded for Illinois, with notes on their distributions Phillips, G. S. 1980. The decapod crustaceans of and habitats within the state. Illinois State Iowa. Iowa Academy of Science Proceedings Academy of Science Transactions 67: 89-104. 87:81-95. . 1975. The life history of the stripetail Prins, R. 1968. Coinparative ecology of the cray- darter, Elheostorna ketinicottt, in Big Creek, fishes Orconectes rusticus and Cambarus tene- Illinois. Illinois Natural History Survey Bio- brosus in Doe Run, Meade County, Kentucky. logical Notes 93. International Revue Gesamten Hydrobiologie 1985. A new crayfish of the genus 53:667-714. Orconectes from the Little Wabash River sys- Pryor, C. W., and C. A. Leone. 1952. Serological tem of Illinois (Decapoda: Cambaridae). Bi- comparisons of astacuran Crustacea. Biological ological Society of Washington Proceedings 103:433-445. 98:564-570. Bulletin , and B. M. Burr. 1973. Distributional Reimer, R. D. 1969. A report on the crawfishes records of the crayfishes Cambarellus puer, (Decapoda, Astacida) of Oklahoma. Oklahoma C. shufeldtii, Procambarus gracilis, P. viae- Academy of Science Proceedings 48: 49-65. viridis, Orconectes lancifer, O. bisectus, and O. , K. Strawn, and a. Dixon. 1974. Notes on rusticus. Kentucky Academy of Science Trans- the river shrimp, Macrobrachium ohione actions 34:51-52. (Smith) 1874, in the Galveston Bay System of Pearse, a. S. 1910. The crawfishes of Michigan. Texas. American Fisheries Society Transac- Michigan State Biological Survey Publication tions 103:120-126. 1:9-22. Rhoades, R. 1944a. The crayfishes of Kentucky, Peck, S. B., and J. J. Lewis. 1977. Zoogeography notes and evolution of the subterranean invertebrate with on variation, distribution, and de- scriptions ol faunas of Illinois and southeastern Missouri. new species and subspecies. Ameri- Natural Speleological Society Bulletin 40:39- can Midland Naturalist 31:111-149. 63. . 1941b. Furthei studies on distribution Penn, G. H. 1942. Observations on the biology and taxonomy of Ohio crayfishes, and the de- scription of a subspecies. of the dwarf crayfish, Cambarellus shufeldtii new Ohio Journal of Science 44:95-99. (Faxon). American Midland Naturalist 28: 644- 647. RiECEL, J. A. 19.59. The systematics and distri- . 1943. A study of the life history of the bution of crayfishes in California. Calilornia Louisiana red-crawfish, C.atnbarus clarkii Fish and Game 45:29-50. Girard. Ecology 24:1-18. -. 1950. The genus Cambarellus in Louisi- RiET/., N. M. 1912. Ecological relations of the ana (Decapoda, Astacidae). American Midland crawfishes of Illinois. B. S. Thesis. Lniversity Naturalist 44:421-426. ol llliiKiis, I'rbana-Champaign. 446 Illinois Natural History Survey Bulletin Vol. 33, Art. 4 .S.viiTH. D. G. 1981. Evidence for hybridization Unger, P. A. 1978. Natural history- invenion of between two crayfish species (Decapoda: Cam- Colorado. 3. The crayfishes (Crustacea: Cam- baridae: Orconectes) with a comment on the baridae) of Clolorado. University of Colorado phenomenon in Cainbarid crayfish. American Museum, Boulder. Naturalist 105:405-407. Midland \'an Deventer. W. C. 1937. Studies on the biology of the crayfish Smith, P. W. 1957. An analysis of post-Wisconsin Cambarus propinquus (Girard). biogeogiaphy of the Prairie Peninsula region Illinois Biological .Monogographs 15(3»:l-67. based on distributional phenomena among \'illalobos, a. 1983. Crayfishes of Mexico (Crus- terrestrial vertebrate populations. Ecology tacea: Decapoda) (Translation by H. H. Hobbs, 38(2): 205-218. Jr.) Baba Barkha Nalh Printers. New Delhi. India). . 1961. The amphibians and reptiles of Illi- nois. Illinois Natuial History Survey Bulletin VioscA, P., Jr. 1931. The bullhead, Ameiurus 28:1-298. melas calulus. as a dominant in small ponds. Copeia 1931:17-19. . 1971. Illinois stieams: a classification based on their fishes and an analysis of factors . 1939. Where to fish in Louisiana for craw- responsible for disappeaiance of native species. fish. Louisiana Conservation Review 8(1):I7- Illinois Natural History Survey Biological 18. Notes 76. . 1953. .\ll about crawfish — life history . 1979. The fishes of Illinois. University of and habits. Louisiana Conservationist 5(6|: 3-5. Illinois Press, I'rbana. Wayne, W. J. 1952. Pleistocene evolution of the Ohio and Wabash \alle\s. Journal of Geology Strenth, N. E. 1976. A review of the systemaiics 60: 575-585. and zoogeography of the freshwater species of Palaeinonele.s Heller of North America (Crus- Weagle, K. \'.. and G. W. Ozbirn. 1972. Obser- tacea: Decapoda). Smithsonian Contributions vations on aspects of the life historv of the cray- to Zoology 228. fish. Orconettes virihs (Hageni, in imnhwest- ern Ontario. Canadian Journal of /oology Tack, P. I. 1941. The life history of the crayfish 50: 366-370. Cambarus imriiunis Hagen." American Midland Webb, D. W., N. D. Penny, and J. C. .Marlin. Natuialist 25:420-446. 1975. The Mecoplera. or scorpionflies. of Il- linois. Illinois Natural Hisiorv Survev Bulletin Teller, T. 1973. Preglacial (Teays) and early J. 31:250-316. glacial drainage in the Cincinnati area, Ohio, Kentucky, and Indiana. Geological Society of White, F. A. 1949. Preliminary notes on the breed- America Bulletin 84(1 1): 3677-3688. ing season of Palaemoneles kndiakcnsis Rath- bun in the Baton Rouge .\rea. Louisiana .Acad- Threinen, C. VV. 1958a. Wisconsin crayfish. emy of Sciences Proceedings 12:71-74. Wisconsin Conservation Bulletin 23(7):13-15. Williams, \. B. 1954. Speciaiion anddisiribution . 1958b. A suininary of observations on the of the crayfishes of the Ozark Plateaus and commercial harvest of crayfisli in northwestern Ouachita Provinces. l'niversit\ of Kansas Sci- Wisconsin with notes on the life history of ence Bulletin 36:803-918. Orconectes viritis. Fish Management Division of the Wisconsin Conservation Deparlmeni , and .\. B. Leonard. 1952. Lhe cia\fishes Miscellaneous Report 2:1-14. of Kansas. l'niversit\ of Kansas Science Bulle- tin 34:961-1012. Truesdale, F. M., and W. Mermilliod. 1979. J. Williamson, E. B. 1907. Notes on the crayfish o{ The river shrimp Macrobrachium ohionc Wells County, Indiana, with the description of (Smith) (Decapoda, Palaemonidae): its abund- a iww species. Pages 749-763 i>i .\nnual upon. ance, reproduction, and growth in the Atcha- Department of Cieologv and N'atur.i! Resources falaya River basin of Louisiana. I'.S.V. Crusta- of Indi.ina. ceana 36: 61-73. Wii.i.MAN, H. B.. aiul J. C. Frvk. 1970. Pleisuxene TtiRNER. C. L. 1926. The ciaylishes of Ohio. Ohio sn.itigiapby of Illinois. Illinois Stale Geologi- State Universilv Bulletin 30:14,5-195. cal Sur\ev Bulletin 94. INDEX •147 ncanthura. Caxibaru.s, 435 wi.scon.siiii'>isis, 335, 417 aumlhurus. Miurnbriu liiuiu. 356 larrinu.'i, Ma< robrat htuiii, 356 aiuliis, C:/unharu\. 376 cau.seyi, Onoiintes, 419 aculus, Cambaru.s btaniiingii. 376 Clarkii, Cambarus. 381 acutus. Pxocambanis, 376 clarkii. Procambarus. 381 alnbametisii. Orconcctcs, 390 Conservation, 440 antrorum. Pnlaemonetes, 359 louesi, Canibarus, 417 argillicota. Canibarus, 335, 422 Crayfish Life Cycle. 343 A.slacus. 422 Creaserinus, 335 jodtens. 422 cuevachicae, Procambarus acutus. 376 fossor. -i'i-i cummingi, Palaemoneles, 359 Barbicambarus, 362 debilis. Canibarus. 417 barloni, Catribarus, 428 Derapoda. 335 Barlonius, 426 diminulus, Cambarellus, 368 biseclus. Orconecles, 338 diogenes, Canibarus, 434 biacki, Cambarellus, 368 Dirigicambarus, 335, 363 blaudingu. Procambarus. 376 Distocambarus, 362 blandingii. Procambarus acutus. 376 Boiu hardma. 362 Erebuurnbarus, 335, 428 brevis, Camharus bartoni. 335, 428 exiUpes, Palaemoneles, 335, 360 Extraliniital .Species and Studies, 336 Cambarellinae, 335 Cambarellus. 335, 363 Fallicambarus. 335, 422 fc/ac*), 368 422 diminutus, 368 fodiens, hedgepethi. 422 /cs/ifj, 368 ulileri, 422 ninae. 368 Faxonelta. 362 pufr, 366 faxonii. Canibarus. 400 schmilti. 368 Faxonius. 386 shufeldtii. 363 immunis pedianus. 388 /cva/H(i, 368 .-istai us, 422 C:ambaridae, 335, 362 fodiens. Fallicambarus. 422 Cambarinae, 335 fodiens. Orconectes, 412 Canibarus. 335, 426 forceps, Astacus. 434 acanthura, 435 fossor, Freshwater Shrimp Life Cycle, 345 af!((uj, 376 argitlicola, 335. 422 bartoni, 428 Geological History of Illinois, 339 ferci'ii. 335, 428 Girardielta. 335, 371 /flpi^w. 428 gracilis, Canibarus, 370 tenebrosus. 428 gracilis, Procambarus, 370 blandingti acutus. 376 C/ar/(H.'381 Historical Studies of Illinois Decapods, 335 couesi, 417 harrisoni. Orconectes, 338. 396 rfc6i7(i. 417 hedgepethi, Fallicambarus, 422 diogenes. 434 Hohbseus, 362 var. Ludoviciana. 434 holthuisi. Palaemoneles, 359 ludovicianus, 435 hubbsi, Canibarus, 338 jaxonii, 400 /iy/as, OrfOfiff(«, 338 gracilis, 370 /H lancifer, Orconectes, 400 kadiakensis, 359 laei'is, Cambarus, 428 paludosus, 358 lan'is, Cambarus bartoni, 428 /pxanui, 359 lancifer, Cambarus. 400 Palaemonidae. 335, 356 lesliei, Cambarellus, 368 paludosus, Palaemonetes, 358 liberorum. Procambarus. 371 Pandicambarus, 335, 368 limosus, Orconectes, 412 Paracambarus, 369 Ludoviciana, Cambarus diogenes var., 434 Pacijasticus. 362 ludovicianus, Cambarus diogenes, 435 pedianus, Faxonius immunis, 388 luteus, Orconectes, 338 peruncus, Orconectes, 338 placidus, Cambarus. 404 placidus, Orconectes, 404 Macrobrachium, 335, 356 Procambarus, 335, 369 acanlhurus, 356 acij/ui, 376 carcinus, 356 acutus, 376 ohione, 356 cuevachicae, 376 oljersii. 356 blandmgii, 376 medius, Orconectes, 338 blandingii. 376. 378 millus, Cambarus, 435 c/arAn, 381 gracilis. 370 r!a«, Orconectes, 338. 419 liberorum, 371 Natural History, 343 reimeri, 371 r!f6raicfn5«, Cambarus, 434 troglodytes, 335 ninae, Cambarellus, 368 inaeviridis, 378 propinquus, Cambarus, 406 propinquus, Orconectes, 406 obesus, Cambarus, 335, 434 pufr. Cambarellus, 366 Macrobrachium, ohione, 356 Puncticambarus, 335 Ohionis, Palaemon, 356 punctimanus, Orconectes, 338 oljersii, Macrobrachium, 356 putnami, Orconectes, 412 Orfonpf/«, 335, 386 alabamensis, 390 quadruncus, Orconectes, 338 bisectus, 338 fOUiiC)!!, 419 forceps, 412 reimeri, Procambarus. 371 harrisoni, 338, 396 rhoadesi. Orconectes. 390 /i5)/a5, 338 robuslus, Cambarus, 439 illinoiensis, 386 ruslicijormis, Cambarus, 432 immunis, 388 ruj()fui. Camfcaru5. 412 immunis x pedianus, 390 ru.s Brown. Britf VV . Ph.D.. Assi.UanI Supportnr S( irnlisl I.F Mfrf. I.IOYD, Teihnnal Hluslrator Dr/A\. Ronald K., Assnlanl Supportn-r Siirntisl Mc;C;LELr.AN, SiHRLFV. B.S., Assof latr Terhnual Editor (.lirri v. David. M.S.. A\si\tanl Supportivr .Snentisl Stfgfr, Eva, B.A.. Assistant Teihnical Editor HbRKtRT. Jamf_s R., M.S., Assistant .Supportivr Siirnltst JacK-SON, Cynthia Ci,. B.A,, A.\si.\lant Supporlnr S< irntisl Technical Library JoNEi. Jon Martin, M.A.. 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(48.816— 3M— 8-85) Some Publications of the ILLINOIS NATURAL HISTORY SURVEY BULLETIN Kankakee River Drainage. Illinois. .Michael J. Sulc and Thomas M. Skelly. 1985. 16 p. Article 1. Dipiera, or True Flies, of Volume 33, — 124. — Illinois Birds: Vireos. Jean W. Graber. Illinois. I. Tabanidae. L. L. Pechuman, Donald Richard R. Graber. and Ethelyn Kirk. 1985. W. Webb, and H. Teskey. April 1983. 122 p. J. 52 p. \'<)lume 33, Article 2.— An Annotated Bibliog- raphy of the Illinois Herpelological Literature 1960-1980, and An Updated Ciheiklist of SPECIAL PUBLICATION .Species of the State. Michael A. Morris, Richard 2. Landsca|)e Ecology: Directions and .S. Funk, and Philip W. .Smith, April 1983. 15 p. Number — Approaches. Paul G. Risser. James R. Karr. Volume 33, Article 3. — 12.') Years of Biological and Richard T. F. F'orman. November 1984. Research 1958-1983: A Symposium. George 18 p. Godfrey, Chairman; John Bouseman, William Edwards. Kenneth Robertson, and Robert CIRCULAR Zewadski, eds., Proceedings of the 125th An- niversary Symposium of the Natural History 49. —The Dunesland Heritage of Illinois. Her- Survey. 1985. 200 p. bert H. Ross. August 1963 (Reprinted .Mav 1974). 28 BIOLOGICAL NOTES p. 51. — Illinois Trees: Selection, Planting, and 120. — Recreational Fishing in the Kankakee Citre. J. Cedric Carter. March 1977 (Third printing). River. Illinois, Robert J. Graham, R. Weldon 123 p. Larimore, and William F. Dimond. June 1984. 52. — F'ertiliiing and Watering Tiees. Dan Neely 13 p. and E. B. Himelick. Decemln-r 1971 (Third 121. — Bibliography of Illinois Vegetation. Paul printing). 20 p. C;. Risser. August 198-1. 51 p. 54. —Corn Rootvvoiin Management in Ciinning 122.—The Life History of the Mud Darter, Sweet Corn. W. H. Luckmann. J. T. Shaw, Elheosloma asjjrigptie, in Lake Greek, Illinois. D. E. Kuhlman. R. Randell. and G. D. LeSar. Kevin S. Gummings, James M. Grady, and March 1975. 10 p. Brooks M. Burr. Decemliei 1984. 16 p. 55. —Observing. Photographing, and Collecting 123.—The Life History of the Shorthead Red- Plants. Kenneth R. Robertson. August 1980. horse. Moxosloma miurolrpidolitin. in the 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 Address orders and corivspondence to (he Chief, Illinois Natural History .Survey Natural Resources Building 607 East Peabody Drive, Champaign, Illinois 61820