The Known Cravfishes of :

A Summarv Renort

Prepared By

Terry Inman Bio.310

For

Dr. Paul C. Marsh ASU Center for Environmental Studies Introduction

The crayfishes ofNorth America display greater diversity in terms of species than

any other part ofthe world. Some 338 recognizedtaxa (308 species and 30 subspecies)

exist within the and Canada (Taylor, C.A. et d,. 1996). At least one species

of crayfish is native to every state in the contiguous United States (Hobbs, H.H. Jr. l9S9)

except Arizona which claims no species of its own. Until recently Arizona had (from our

earliest records) remained void ofthis diverse group of invertebrates. However, sometime

within the last 30 years crayfishes have begun to show up in Arizona waters. Some have

been intentionally stocked by the Arizona Game and Fish department (AZGFD stocking

records l93l - l99l) and the U.S. Fish and Wildlife Service (verbal - MarstU P.C.) as

forage for game fish such as trout and large mouth bass. Although the extent ofthe

introductions by the USFWS in not known, the AZGFD has reported three sites of

introduction (all lakes), one inl97l, and two in 1991. other sources of crayfish

introduction have probably been a result of bait bucket introductions by sport fishermen.

Although no direct evidence ofthis is known it is perhaps the most reasonable explanation

for their wide spread existence.

Despite crayfishes having become common fauna in many Arizona waters they have managed to attract little attention from any Federal, Statg or scientific community.

The kinds, distribution, and abundance of Arizona's crayfish is unknown, they have not yet been inventoried, or studied. Dr. Paul Marsh from Arizona State University's Center for Environmental Studies was among the first to recognize the need for an in depth look at crayfish in Arizona. Under the direction of Dr. Marsh it has been the scope of my study to acquire crayfishes from around Arizona (by collection and receiving from other sources), to note where they were obtained from, to identify each individual to species if

possible, to catalog each collection, and prepare a distribution map and a sunmary report

of my findings.

Collection

Crayfish may be collected by a variety of methods such as seining (using aYq inch

mesh), minnow traps baited with meat, dip netting, and electrofishing, all of these methods

have proven effective. While dip netting, electrofishing and seining are all active methods

of capture, baited minnow traps are not, and may require an overnight stay in order to

yield the best results. The type of bait used may also play a significant role in the success

of collection efforts. My experience and the findings ofKutk4 F.J. et aL. (1992) suggests

that fish pieces are a highly attractive bait to crayfish, and oily fish (such as smelt, or

anchovies) are most often preferred to non oily fish (such as cod and pollock).

Crayfish are best preserved by placing them in a solution of 6 percent neutral

formalin forlZ hours or up to one week depending on the size (Flobbs, H.H. Jr. 1972).

After being rinsed in running water for a few hours, specimens should then be transferred to a70 percent ethyl alcohol solution, o4 a20 to 30 percent isopropyl alcohol may also be used (flobbs, H.H. Jr. 1972).

Identification

In order to accurately identify crayfish to species it is necessary to have a'form f' male. From I males (or sometimes referred to as "lo form") occur in the subfamilies

Cambarinae and Cambarellinae. Males in both subfamilies undergo cyclic dimorphism that is associated their reproductive cycle. Crayfishes of more northern climates follow a

circular rhytfun, that is to say that at the end of their first season breeding males molt and

are returned to a juvenile morphology (known as'form If" or'2d form') until the next

breeding season when the semiannual molt returns them to form I breeding males. This

cyclic regression takes place throughout the life span of breeding males. It is typical in

Northern populations to find all adult males in either the form I or form II state. However

in species occurring at lower latitudes there is no well defined breeding season, therefore,

form I and form II males may occur in many populations at the same time (Hobbs, H.H.

Jr. 1972). Form I males maybe distinguished from form II and juvenile males by the

occulrence of"corneus, or horney, terminal elements (projections) on the distal ends of

the first pleopods" (rlobbs, H.H. Jr. 1972) (see appendix a, fig 2 for pleopods). It is

important to note that the size ofthe crayfish will not determine whether it is in form I or

form II. Molting increases the size of the crayfish and molting rnay also regress the

individual to a form II state, because of this, a form tr male may be larger than a form I

male. Most identification keys are based on form I males since their pleopod morphology

is much more distinct from the pleopods of form fI males and the juveniles of other closely related species. In North America the usage of from I males for identification applies to

all crayfishes except for those belonging to the genus Pacifastacus which do not exhibit cyclic dimorphism. The following appendices relate to the anatomy, identification and catalog specimens ofArizona's crayfish species: appendices d B, and C. Arizona's known crayfish species

I have separated Arizona's known crayfish species into two categories; the first

category is the crayfishes that are known to occur in wild populations, and the second

category is the crayfishes that are known to occur in pet stores and therefore private

aquariums. It is important to mention that the collection efforts are still on going and that

not every region ofthe State has been surveyed or exhausted in collection efforts, hence

the usage of the words "known crayfishes".

The wild crayfishes that have been collected thus far are Procambarus ctarkii

whose cofilmon name is the'?ed swamp crayfish" and Orconectes virilis whose common

name is the'horthern crayfish". Proccnnbarus clarkii has a native range from north-

eastern Mexico and south-central USA west to Texas and east to , north to

Tennessee and (floldich and Lowery l9s8). It is among the most widely

distributed of all crayfish species in the United States and has been successfully introduced

into fourteen countries worldwide (Holdich and Lowery 1988). Despite the apparent

popularity af Procarnbarus clarkii as a food source in the southern United States, it is

considered a pest in some countries (as well as some areas in the United States) such as

Japan where it is blamed for the destruction of Rice seedlings, taro roots, and fishing nets.

The often intense burrowing activity by large populations of Pracanbarus ctarkii has also been known to cause extensive damage to earthen dikes and other water control structures

(floldich and Lowery l98s). The habitat preferences of P-. clsrkii include lentic and lotic habitats, with fine silt to gravel substrates. They are also known to burrow (see appendix

D for an Arizona distribution map). Orconectes virilis is the most widely distributed Canadian species of crayfish with

a native range that spans from Saskatchewan to Ontario, eastward to , and from

Montana to and east to . Orconectes virilis has been introduced into several

states from the east coast to the west coast. While it has not achieved the popularity of^P.

clarkii as a human food source (most likely due to its smaller size), it has achieved

popularity as a'bait fish", this may in part explain its substantially expanded range. The

habitat preferences of Orconectes virilis ranges from lentic to lotic waters with substrates

that range from silt to cobble (see appendix D for an Arizona distribution map).

The pet shop varieties ofcrayfishes that I have collected thus far have been

Procambarus alleni and one advertised tobe Procambarus paeninsulanus. The

Procambarus alleni specimen came from Pets Inc. a small pet store on the north-west

corner of Nfill and Southern in Tempe. Pets inc. sells them as forage for oscars and other

large aquarium fishes. Upon inspection ofthe aquarium small white translucent egg

shaped sacks could be seen on the thorax of each crayfish, some with many (ca. 20 -30)

and others with just a few (5-6). The specimen that I procured only had a few of these

egg like sacks, and upon preservation ruptured. No identification ofthe sacks has yet been made. The stores owner is aware of the sacks, but does not know what they are, just that'they come this way''. No name was given on the tank and the owner nor his employees knew the genus, species or common name of the crayfish they carry.

Procambarus alleni is not a widely distributed crayfish species, their entire range is within the State ofFlorida where their numbers are currently stable. They occur in both lentic and lotic environments and prefer to burrow. Petco at the south-east corner ofMcClintock and Warner Roads in Tempe advertise their crayfishas Procambwus Weninsulanus (which were also being sold as forage and as a unique addition to freshwater aquariums), however, upon identification it turned out to be Procambarus clarkii. This miss representation carries many implications, for example it may mean that the pet store doesn't know the species it receives, and perhaps the distributor doesn't know what species it is distributing. However, it should not be assumed that Procambarus paeninffilarrus has never been canied or that it won't be carried at that pet store in the fufure. Procambarus paeninsulamts has a native range of

Alabama, Florid4 and and has not been introduced elsewhere. The habitat preferences of Procambarus Weninsulanus is in both lentic and lotic situations where it may construct burrows.

There are 13 species and 3 subspecies of crayfish that neighbor Arizona in the

United States, however only two species ?. clarkii, and O. virilis have been found to border Arizona. Mexico has 44 species and 2 subspecies of crayfish within its boundaries, however, to date none ofthem border the State of Arizona. Appendix E has a complete crayfish species list for each state that surrounds Arizona as well as a species list for

Mexico; appendix E also contains a distribution map for those species that border Arizona.

Conclusion

Arizona is unique to the rest of the contiguous United States and most ofNorth

America in the factthat it has no native crayfish species of its own. However, with the relatively recent introduction and establishment of8. clarkii and O. virilis it apparent that they have taken up permanent residence in Arizona. It is also obvious to see that the door is open for the introduction and possible establishment of other crayfish species such as 8. alleni and 4. Weninsulanus (as well as a potential parasite) from private aquarium introductions. The only crayfishes not allowed by law in Arizona are the Australian crayfishes from the family Parastacidag therefore, the door for "domestid' (those found in the United States) crayfish introductions is seemingly wide open.

There is also a need for future research to examine the effects of L. clarkii and O. virilis on our aquatic ecosystems and the native plants and animals that inhabit them. I. clarHi is already known to have disastrous effects on some ecosystems, destroying earthen water cnntrol structures and commercial crops (floldich and Lowery 1988). O. virilis has been implicated in causing the decline of lake trout populations in the Great

Lakes (Savino and Miller 1991) as well as causing the trout fishery at Utah's New Castle

Reservoir to crash (Hepworth and Duffield). Certainly two crayfish species with the potential to caus€ such damage in an ecosystem warrants the need for future investigations, especially when considering Arizona's fragile native fish populations. BibliosraBhy

Arizona Game and Fish Department. 1931 - 1991. State wide stocking records

Hepwortll D.K. and D.J. Duffield. 1897. Interactions between an exotic crayfish and stocked rainbow trout in New Castle Reservoir, Utah. North American Journal of Fisheries Management. Volume 7. 544-561.

Hobbs, H.H. Jr. 1972.Biotaof Freshwater Ecosystems. Identification Manual No. 9.; Crayfishes (Astacidae) ofNorth and Middle America. Environmental Protection Agency Project # 18050 ELD, Contract # 14'12-894- U.S. Government Printing Office. , D.C.

Hobbs, H. H. Jr. 1989. An illustrated Checklist of the American Crayfishes ( Decapoda: Astacidae, Cambaridae, and Parastacidae) Smithsonian Contributions to Zoology. Number 480. Smithsonian Institution Press. Washington, D.C.

Holdich, D.M. and R.S. Lowery 1988. Freshwater crayfish: biology, management, and exploitation. Croom Helm Ltd. London, England.

Kutk4 F.J. et aL. 1992. Bait preferences and trapability of two common crayfishes in Northern Minnesota. The Progressive Fish-Culturist. Volume 54:254-254.

Savino, J.F. and J.E. Mller. 1991. Crayfish(Qreones!9fffiAs) feeding on young lake trout (Satveluru;nmrqvcash): efect of rock size. Journal ofFreshwater Ecology. Volume 6, Number 2 - June,1991.

Taylor, C.A.; et al. 1996. Conservation status of Crayfishes of the United States and Canada. Fisheries Vol. 2l(a). American Fisheries Society. Bethesd4 MD. Cravfish Anatomv

AnoendixA

Taken from Hotibe, H.H. Jr. Id manual # 9 (lW2) suborbital angle

vical spine length of carapace

width of areola length of areola areola

Fig. 1. Dorsal view of generalized male crayfish iLLustrating structures and neasurements referred to in keys. width of oalm

propodus of chela length of palm

antenna antennal scale antennule

ischium of 3rd maxilliped

coxa of chelioed

hook on ischium

boss on coxa

pleopods I

Fig. 2. Ventral view of generalized male crayfish illustrating stnrctures and measurements referred to in keys.

t f2- re nt ra I Proi ect io n --5-1 ( tC-caudal knob\-) \ proces'Xlj L__fmesiar ffi_spermsroove_p /ffi-spermsrooven ( t lt\ n

Fig. 3. First pleopods of nenbers of Canbarinae and Canbarellinae. 4,b, Mesial and lateral views of terminal portions of generalized pLeopods illustrating arrangenents of terninal elenents; c, Methods of measuring first pleopods (mesial view) in 1reonectes (e, central projection; nP, mesial process); d-i, Lateral views of left pleopods of first and second form na1es, respectively, with corneous central projection shaded - d, ConbatelLus; e , Canbanr,s; f , )reonectes; g, Procottbarus i h, Hobbseus; i, Eallicanbtus); j , Caudal view of left pl.eopods of first and second forn nales, respectiveLy, Ln Fa,ronelLa (corneous central projection shaded).

10 Kev to Arizona's Cralfish Snecies

Anoendix B

Taken from Hobbe, H.H. Jr. Id manual # 9 (1972) SECTION II

KEY TO THE MALE MEMBERS OF THE NORTH AIUERICAN CMYFISH GENERA (Except for Paeifastacus, a1l based on First-Form Male)

NOTE: Before beginning to use this key, it witl ue h6rprul to de- terrnine whether the first pleopods are synnetrically or asynnetric- a1ly arranged--see Figure 2L.

1 Ischia of all pereiopods lacking hooks (Fie. 4f; see also Fig. 5a): RSTACINAE: Paeifastaeus Bott, 1950 (p, 2L) Ischia of I or nore pairs of pereiopods with hooks (Fig. 4a-c)

/\ ./\/< ,/ \t ./l /t A /\, ./l fi'g\z €'4i\ P<-/, ft71 {&, rgdt A L.(t/ ) '/913' P [z/rt/) \,€flt^ ,ffi w\&.,tj'} \ .--1 ,/ \ KW' (&-/ (s \fr/ v{> w tr >tfu, "av" " ^v ,w \-/ ,w .w\-/ 's)-

Fig. 4. ventral view of basal portions of left pereiopods with ischia bearing hooks shaded.

2(L) rschia of second and third pairs of pereiopods with hooks (Fig. 4b; see atso Fig. 5c): CAagAnrLLtNer: CottbatelLas 0rtnann, 1905a (p. 25) rschia of second pair of pereiopods. never bearing hooks- (Fig. 4a,c-e): CAMBMINAE...... 3

3(2) Body pigrnented; eyes faceted and pignented 4 Body albinistic; eyes seldom faceted and with pignent greatly reduced or absent ....L2 4 (3) First pleopod terninating in 2 elenents; long, slender central projection; short nesial process no nore than 1/2 length of central projection (Fig. 5w); central projections of paired pleopods overlapping (Fig. 19): FasoneLla Creaser, 1933 (p. 29) First pleopod terminating in 2 or more el.ements; if with 2, nesial process nore than L/2 Length of central projection; central projections of paired pleopods never overlapping (Fig. Sd-v) except occasional.l.y in Fallicqnbamrc dissittts in which also recunred at nore than 90 degree angle (Figs 81c,82d)b.... 5

s (4) First pleopod terminating in 2 elenents (Fig. sh-i, 1-p).... 6 First pleopod terninating in more than 2 elements (Fig. 5d-g, k,Q,v) 10

6 (s) Distal L/3 of first pleopod with prominent shoulder on cephal- ic surface (Fig. 5h), or" central projection forming dis- taLLy projecting triangular plate (Fig. 5j), or central projection arising from enlarged subterminal or tenninal area (Fig. 5i) i Proeantbamts 0rtmann, 1905b (part) (p. 31) Distal 1/3 of first pleopod lacking shoulder on cephalic sur- face; central projection never foming distally projecting triangular plate nor arising fron enlarged subterminal or terminal area (Fig. 51-p, r-u) 7

7 (6) Coxa of fourth pereiopod lacking caudolnesial. boss (Fig. 6a)... 8 Coxa of fourth pereiopod with caudonesial. boss (Fig. 6b-d).... 9

Fig. 5. a-v, Lateral view of left first pleopodsi w, Caudal view of same. a, Pacifastaclls L. troubridgii; b, Trogloeonbants macLqneii c, CqnbayelLus pueri d, Pnocanbants fitzpatz.iekii e, Proeqnbants paeninsulqrtttsi f , Pyocqnbwus a. aeutus, g, pz,o- corbants aduena; h, Pnocattbayus meui.eqnus! i, Proeqnbatus tm,rculentus; j, Procantbarus pAgnaeusi k, proeanbarus pffiadorus; I, 0reonectes Limosus; m, 2rconectes propinqttusi tr, 2neoneetes innnutisi o, Hobbseus cristatusi p, Hobbseus oreoneetoides; Q, Cmbants pristinusi r, Canbuus b. baytoniii s, Conbamrc reduneus; t, FaLLieanbarus hortonii u, Fallieanbants fodiens; v, Fallieqnbarus maeneeseii w, Fa,xonelLa ereasert.

15 i

8 (7) First pleopods, in resting position, deeply withdrawn between basLs ol pereiopods and largely concealed by dense setifer- rous nat extending fron ventrolateral nargins of sternun (Fig. 7a): Hobbseus Fitzpatrick and Payne, 1968 (p. 75) First pleopods, in resting position, never deeply withdrawn between- bases of pereiopods and never concealed by dense setiferous rnat extending from ventrolateral nargins of sternur (Fig. 7b): Oreoneetes Cope, L872 (part) (p. 77)

il li Fig. 6. Ventral view of basal podomeres of left third, fourth, and fifth pereiopods showing variations in coxae (stippled) of fourth t, (a, without boss; b-d, with boss) . 4, Procqmbavws digaeti; b, P. tit paeni,nsuLanus; c, P, r'Lojaii d, P. tenuis. i j 9(7) Opposable nargin of dactyl of chela with abrupt excision in proxinal 1/2 (Fig. 8a): EaLlicanbatus Hobbs, 1969b (part) (p.ee) in _Opposable nargin of dactyl of chela without abrupt excision - proxinal L/2 (Fig. 8b-e): Cqnbazas Erichson, 1846 (part) (p.103)

Fig. 7. Ventral thoracic region of a, Hobbseuei b, )z.eoneetes.

l6 10(5) Central projection of first pleopod bladelike, al,ways directed caudally or caudodistally, and bearing subterminal notch (n) (Fie. 5q) : Cqnbartts (part) (p. 103) Central projection of first pleopod seldon bladelike, if so, directed laterodistaLly or lacking subterminaL notch (Fig. Sd-g, v).. . tl

Fig. 8. Dorsal view of right chelae. &, FalLieqnbants oryktes; b, Conbutts Latimattus; c, C. Longalus; d, C. eatraneusi et C. braehydaetylus,

11 (10) First pleopod with central projection and mesial process curved caudally at at least 90 degrees to principal axis of shaft of appendage (Fig. 5v): Fallicottbuas (part) (p. 99) , First pleopod never with both central projection and nesial process curved caudally at angle of so nuch as 90 degrees to principal axis of shaft of appendage (Fig. 5d-g): Proeonbants (part) (p. jt)

Fig. 9. Ventral view of left third naxillipeds. a, Troglo- eqnbarzts maelqneii b, Proeqnbarus pallidus. (1, ischiun. )

L7 KEY TO SUBGENEM OF PROCA]LBARUS (Based on First-Form Male)

Carapace with 2 or nore cervical spines (Fig. 20a), 2 Carapace with or without 1 cervical spine (Fig. 20b). 4

Fig.20. Lateral view of carapaces showing presence (a) and absence (b) of cenrical spines.

2(L) First pLeopods asynnetrical (Like Fig. 2Lc,d) z Pennides Hobbs, L972 (p. 67) First pleopods syrnmetrical (Like Fig.2La,b) 3

Fig. 21. Ventral view of first pleopods: a and b, symnetrical; c and d, asymmetrical. &, Proeanbayus Vwgeniari,usi b, p. Llanasi; c, P. seminolaei d, P, aeuti.ssirmts. (r, proximonedian lobe.) 3(2) Albinistic; eyes reduced and without pignent: Remotieonbamts Hobbs, 1972 Monotypic : Proeanbatus (n. ) peeki Hobbs , L967. (Subterranean waters of northern Alabama. See Fig. zZb. Literature: Hobbs , 1967) Pignented; eyes well developed: Austy,ocanbarus Hobbs, L972 (part) (p.38)

3T 4 (1) Cephalic surface of first pleopod with prominent angular or sub- angular shoulder situated far proximal to base of terninal elenents (Figs 2Zas,b,c, 30, 56) .. 5 Cephalic surface of first pleopod with or without prominent angular or subangular shoulder; if present, situated irunedi- ately proxinal to base of terninal elenents (Fig. 22d-h), o!, if situated nore proxinally, somewhat rounded and cephalic process never broadly rounded (Figs 48f, 49h, 54a,b) 7 s (4) Hooks on ischia of third and fourth pereiopods (Fig. 4c); first pleopod with subapical setae #,:ttt;#;)rzs Hobbs , rs72 (p. 7r) Hooks on ischia of third pereiopods onLy (Fig. 4a); first pleopod without subapical setae (Fig. 22b,c) 6

Fig. 22. Lateral view of Left first pleopods. a, Procanbaz'us okaloosaei b, P. peckii c, P. rodriguezi; d, P. fitzpatricki; a, P. &Lgueti', f , P. LatipLeumtni E, P. s. sirruLcnsi h, P. tenuis. (s, shoulder. ) 6(5) Mesial process of first pleopod massive, obscuring entire renainder of distal portion of appendage in caudal aspect (Fig. 23a; see also Figs 1lg, 22b) z Remotieqnbamts itobbs, 1972 Monotypic : Pnoeonbwus (R. ) peeki Hobbs , L967. (See couplet 3 for range and literature) Mesial process of first pleopod spiniform or lanceolate, never obscuring entire remainder of distal portion of appendage in caudal aspect (Fig. 23b)z Austrocatnbamts (part) (p. 38)

7 (4) Hooks on ischia of fourth pereiopods only (Fig. 4e), occasion- ally with vestigial ones on third pereiopods...... 8 Hooks on ischia of third or third and fourth pereiopods well developed (Fig.4a,c,d) L0

32 e (8) Mesial ranus of uropod with distonedian spine projecting beyond margin oframus (Like Fig. 40b; see also Fig. 24d): Acuearda Hobbs, L972 Monotypicz Proeanbarus (A.) fitzpatrieki Hobbs, L97I. (Burrows in southern between the Wolf and Pascagoula rivers. Literature : Hobbs, 1.971a) Mesial ramus of uropod with distornedian spine never projecting beyond rnargin of ramus (Like Fig. 40a): (P. ViLlaloboszs Hobbs ' 1972 73) 10 (7) Chel.a without tubercles or brush of setae on mesial surface of palm; first 3 pairs of pereiopods with conspicuous brush of plumose setae extending fron basis to at least proximal part of nenrs (Fig. 25a): CapiLLi.eanbazzrs Hobbs, 1972 (p. 44) Chela with tubercles (Fig. 26a,c-e) (sornetines obscured by conspicuous brush of setae, Fig. 26f) on rnesial surface of paln (tubercles alnost obsolete in P. youngi); first 3 pairs of pereiopods always lacking conspicuous brush of plunose setae extending from basis to at Least proxirnal part of nenrs (Fig. 25b). 11

Fig. 25. Ventral view of Left third naxillipeds and first three pereiopods. a, Procattbarus hinei; b, P. barbakts.

11 (10) Chela strongly depressed, usually broadly triangular, and with nesialnost row of tubercles on paln cristiform or subcristiforn (Fig. 26a) : Hagenides Hobbs, L972 (p. 48) Chela subovate to cylindrical, nostly elongate, ild lacking cristifonn row of tubercl.es on nesial nargin of palm (Figs 26b-f, 38a-c) 12

34 12(11) Hooks on ischia of third pereiopods only (Fig. 4a).. 13 Hooks on ischia of third and fourth pereiopods (Fig. 4c,d).. 16 L3(L2) First pleopod wirh subapical setae (Fig. 22f): Lecontieambamts Hobbs, 1972 (part) (p. Sl) First pleopod without subapical setae (f igi Z2e ,g,-'27ai . .': . . L4

Fig. 26. Dorsal view_of chelae. 4, proeattbarus togersi eryLehts; b, P. youngii e, P. Leeonteii d, p. hirsutust e;p. sewinoLae;- f , P. baxbatus,

14 (13) cephalic process of first preopod forming curved plate across cephalodistal extrenity of appendage (Fig. 27a): MeytcanbarLts Hobbs , Lg72 Monotypicz Proeonbarus (M.) bouuieri (Ortmann, lg09). (Streams in the vicinity of Uruapan, Michoac6n, M6xiio. Literature: Villalobos, 19SS) cephalic process of first pleopod variable in fo::n but neveri consisting of curved prate across cephalodistal extrenity of appendage (Fig. 22e,g) ...15 15(14) Rostrrmr with nedian carina (Fig. 28ak); first pleopod with proxinomedian lobe absent (Like Fig. 2lb; see also Figs 6a, 22e): Proeqnbwus Ortnann, 1905 Monotypicz Proeanbarus (p.) dtgueti (Bouvier, fSgT). (streans in Jalisco and Michoac6n, M6xico. Literature: Villalobos, 1955) Rostrum without nedian carina (Like Fig. 2gb); first pleopod with proxinonedian Lobe very strongly developed (rig. 2tan1: Girardiella LyLe, 1938 (p. 45)

35 Key to Species of Subgenus Leeontieonbans Albinistic; eyes with reduced pignent (See also Fig. 11f): Proeonban)s (L.) mi,LLerL Hobbs, L97L (Subterranean waters in Dade County, Florida. Literature: Hobbs, 197lc) Pignented; eyes norrnally pignented 2

2(r) Hooks on ischia of third pereiopods only (Fig. 4a).. 3 Hooks on ischia of third and fourth pereiopods (Fig. 4c)..,., 6 3(2) Mesial process (tn) of first pLeopod nassive and subspatulate (Fig. 43f): Proeanbarus (L.) kilbai, (Hobbs, 1940) (Burrows and lentic habitats in Calhoun and Gulf counties east and south to Levy County, Florida. Literature: Hobbs, L942b) MesiaL process of first pleopod slender and tapering to acute or subacute apex (Figs 4Ta-e,g,h, 44a-d) ... 4

Fig. 43. Lateral view of left first pleopods. a, Procanbarus VwbbeLli; b, P. natTtbnnaei c, P. capilLahts; d, P. sherrnani; a, P. alleni; f , P. kiLbAi; E, P. Latipleumtrn. h, P. batbatus. (e, caudal. process i fr, mesial process.) 4(3) First pleopod with mesial process extending no farther distally than cephalic process; subapical setae in apical cluster (Fig.43a): Proconborus (I.) hubbeLLi, (Hobbs, 1.940) (Burrows and lentic habitats in the Choctawhatchee River basin in Alabana and Florida. Literature: Hobbs, 1942b) First pleopod with mesial process extending nuch farther distally than cephalic process; subapicaL setae arranged in linear series on cephalodistal nargin of appendage (Fig. 43b,c) ...... 5

51 5(4) First pleopod with caudal process (c) directed cephalodistally, ,'""if,ing level of tips of cephal.ic process and central projection"ot (fig. +sU; see also Fig' 28b): Proeanbants (i. ) rathbunae (Hobbs ' 1940) (Burrows and lentic habitats in okal-oosa and Holnes counties, Florida. Literature: Hobbs, L942b) First pf"op6a with caudaL process- (g) directed distal'ly and extending at least to llvel of tips of cephalic process and central Projection (Fig. 43c)z pioeanbutts (I. ) eapiLlafzrs Hobbs , I97L (Burrows and lentic habitats in the Escambia River basin in Alabana and Florida. Literature: Hobbs, 1971b)

6(2) First pleopod with nesial process (nr) reaching no farther distally than tip of cephatic process (Fig' 43d): - Pv'oeanbwtts (L.) sherTnani Hobbs, 1942 (Strearns and sloughs in Pearl River County, Mississippi' st. i"rr.ny parish, Louisiana, and in the Escanbia River drainage, Florida. Literature: Hobbs, L942b) First pfelpoa with mesia1 process reaching nuch farther distally tt tip of cephalic process (Figs 43e-h , 44a-d) 7 "i 43e): 7 (6) First pleopod with rnesial process sinuous (Fig. Proeonbeus (I. ) aLLeni (Faxon, 1884) (Lentic and Lotic habitats east of the St. Johnrs River and all of peninsul.ar Florida south of Levy and Marion counties. Literature: Hobbs , L942b) First pleopod with nesial process straight or curved but never sinuous- (Figs 43f-h, 44a-d) and 8 (7) First pleopod with mesial process massive, subspatulate, its' apei directed "t^.'f,,;z*i:H,,iil; ilil;, (Hobbs, r e40) (See coupl"et 3 for range and literature) firit pteopba with rnesial process acute to subspiculiform, and its^apei directed caudo- or cephaLodistally (Figs 439,h, 44a-d) or e (8) First pleopod with rnesial process directed distally cephalodistallY (Figs 43g,h, 44a)..,.. 10 First pleopod with nesial Drocess directed caudodistal lY (Fie.44b-d) L2 Fig 10 (e) Palrn of chela of nale not bearded (Fig ' 44e; see also 43g): proeanbants (Il) LatipLeururn Hobbs, 1942 (girrows and lentic habitats in Gulf County, FLorida. Literature : Hobbs, 1942b) Palrn of chela of male bearded (Fig. 44f)' ""'11

52 Key to Species of Subgenus Seapulicambons

Cephalic process of first pleopod acute (Fig. 56a,b) 2 Cephalic process of first pleopod lobifonn with or without angle on caudal nargin (Fig. 56c-e) 3 2(L) Dista1 portion of first pleopod tapering distalLy, in Lateral aspect, fron level of shoulder (Fig. 56a): Proeqnbmus (5.) houellae Hobbs, 1952 (Lentic and lotic habitats in the lower piednont and upper coastal pLain of Georgia between the Oconee and Ogeechee tivers. Literature: Hobbs, 1952b, L962) Distal. portion of first pleopod not markedly tapering distally, in lateral aspect, fron 1evel of shoulder (Fig. 56b; see also Fig. 6b): Proeanbamts (5.) paeninsularuts (Faxon, 1914) (Lentic and lotic habitats fron the. Choctawhatchee drainage in Florida to the St. Johns and St. Marys rivers, and from the Hil.l.sborough River, Florida, northward to southeastern Alabarna and southern Georgia. Literature: Hobbs, L942b, Le62)

Fig. 56. Lateral view of left first pleopods. a, Pz,oeonbayus houeLlaei b, P. paeninsulqnusi c, P. okaLoosae; d, P. cLatkii; a, P. troglodytes. (q, cephalic process.) 3(1) Areola less than 12 times longer than broad (see also Fig. 56c) : Proeanbuas (,S. ) ol

7L i,. l, f 4 (5) Caudal nargin of cephal.ic process of first pleopod with distinct ti (,S. I angJ.e (Fig. 56d): tuoeqnbants ) elatkii (Girard, 1852) ll (Lentic and lotic habitats fron southern Illinois to northern !' I M6xico and Escambia County, Florida. Introduced in Cali- , and Penn, 1943; Hobbs, 3t fornia, Virginia. Literature: RL 1962) s Entire nargin of cephalic process of first pleopod rounded, $r f; lacklng angle (Fig. 56e): fil Proeanbants (5.) trogLodytes (LeConte, 1856) fl [, (Lentic and lotic habitats in the lower piednont and coastal fi,i i!il pl.ain fron the Pee Dee drainage system in to flr the Altamaha drainage in Georgia. Literature: Faxon' 1885; ii'i Hobbs, L962) ti,' ;il i$lr . ili; flil iilil ti lli li;i iii; iii

I lrl rl

il it iir li,i ii:i i

72 KEY TO SPECIES OF GENUS ORCONECTES (Based on First-Form Male)

NOTE: In using this key, the first pleopod nust be viewed nesiaLl.y. For comparative purposes, aL1 of the illustrations of this appendage are made of the left nernber of the pair. see Fig. 5c for tenrinoLogy and nethods of measurements. 1 Central projection of first pleopod constituting L/4 or less of total length of appendage (Figs 60, 63, 65, 67c-g) 2 Central projection of first pleopod constituting nore than 1/4 totaL length of appendage (Figs 68c-g,70, 72,74,77)...32

2(L) Albinistic; eyes without pigurent or facets s Pignented; eyes with pigment and facets 8

Fig. 60. Mesial view of left first pleopods. t, 2yeonectes pellueidus; b, 0. ineomptus; c, 0. a. ausLzalisi d, 0. a, paekwdi; e, 0. i. inerrnis; f , 0. n. fustii; g, 0. Ltstyisoni. (s, shoul.der. ) 3(2) Mesial process of first pleopod extending distinctl.y farther distall.y than central projection (fig. 60a; see also Fig. 11b): Oreoneetes peLLuatdus (Tellkanpf, 1843) (Subterranean waters from Hart County to Trigg County, Kentucky, and Montgomery County, . Literature: Hobbs and Barr, L972) MesiaL process of first pleopod extending only slightly, if at iii: ::::::: :i:::iI :i:l :::::ii i::::::i:: ::::: l?l:il ; i .r:r

:.t:. r:':

i:*t Ai ,lFr'

Si 1::g. -4. 77 9: *. '.' & trS, f,*. G' 32(L) Hooks on ischia of third and fourth pereiopods (Fig. 4c); pLeopod broadened and deeply lii nesial. Process of first grooved distalLY (Fie. 68c): Zreoneetes perunans (Creaser, fg3f) tit (streams in the headwaters of the st. Francis River in lrl 1934a; tri Missouri. Literature: creaser, I:i southeastern lrl Willians,1954a) Hooks on ischia of third pereiopods onl.y (Fig. 4a) , excePt in :{ iil rnesial ; rl,l sone popuLations of 0. hYlas; if present on fourth, tx' I (Figs 68d-g 70, t:if process never narkedlY broadened distally , tt 33 'l,l 72, 74, 77).. r{,, I

1 .a ,fii

'd4 g fr i{ji

iilr1

I{i1 ini left .l, Fig.68. &,b, Dorsal view of carapaces; c-g, Mesial view of a, 0Teonectes LepVogonopodus; b, spinos-us; first pleoPods. 'e, ,0' 0' c, 0. peruncus; d', 0, putnuni; b- Leptogonopodus; f ' medius; E, 0. spinosus. 33(32) Central projection of first pleopod_constituting at least L/2 total length of appendage- (Fig' 68d-g) "':' li. Centrat ptoiE"iiol of firsi pieoioa conitituting less t'han I/2 total length of appendage (Figs 70, 72, 74, 77) 37 34(33) Length of nesial nargin of palm of chela no more than 1/2 length of dactyl (pig. Oga; see aLso Fig' 68d):- lreoneetes Pitrnord (Faxon, 1844) (streams in the drainage in western Kentucky, southern (?),andTennessee(?).Thelinitsoftheranges of thii rp""i"t and the closely allied 0, iuueni,Li's and 0. fiiii spinosus irave not been deterrnined. Literature: Ortnann, lg5l,treatedthisspeciesando.spinosusassynonymsof 0. jut;eniLi's. Length of nesial nargin of Paln of chela greater than L/2 length of dactYl (Fig. 69b-d) {llll

ffilli 86

ilil. Fig. 69. Dorsal view of chelae. a, 1yeoneetes putnnni; b, 0. Leptogonopodusi c, 0. mediusi d, 0. spinosus. 35(34) Rostrrrn with median carina (Fig. 68a; see also Figs 68e, 69b); 1reoneetes Leptogonopodas Hobbs, 1948 (Streams in the Red River drainage system in Arkansas and easterrl . Literature: Fitzpatrick, 1965; Wil1ians, 19 54a) Rostnrn without nedian carina (Fig. 68b). 36

36(55) Chela with large proninent punctations dorsally; width of pahn greater than 1.4 tines length of its nesial nargin (Fig. 69c; see also Fig. 68f): )reoneetes medius (Faxon, 1885) (Streans in the headwaters of the Big and Merarnec rivers in southeastern Missouri. Literature: Creaser, 1934a; Wil.Lians, 1954a) Chela with fine punctations dorsal.ly; width of paLn less than 1.4 tines length of its rnesial nargin (Fig. 69d; see also Fig. 68b,g): )rconeetes spinosus (Bundy, L877) (Strean tributaries of the Coosa, Kanawha (?), and Tennessee rivers in ALabana, Georgia, eastern Tennessee, Virginia (?), and (?); see statenent in couplet 34. Literature: Ortmann, 1931, treated this species and 0. putnoni as synonyms of 0. juueni,Lis.)

37 (33) Both terminal elenents of first pleopod curved caudally or caudodistally (Figs 70, 72).. 38 Mesial process of first pleopod never directed caudally or caudodistally (Figs 74, 77).. 52

87 Fig. 70. Mesial view of left first pleopods. 4, )teoneetes hobbsi! b, O. p. palmerli c, 0. p. ereoLqttusi d, 0. P. Lorqi,marus; a, 0. eotrPressus; f , 0. aLabanensis; E, 0. quadrunct'ts. along part of length or so reduced in width 38(37) Areola obliterated (Fig' as to accomrnodate no punctations in narrowest part 71a,b) 39 Areola broad or narrow but always with roorn for at least one punctation in narrowest part (Fig' 71c,d) 42 39(38) Areola not obliterated along any part of length,-(Fig. 71a; see alsoFig.70a):7reoneeteshobbsiPenn,l9S0 (Strearnl in tire Lake Pontchartrain watershed in Louisiana and Mississippi. Literature: Penn, 1952) Areola obl.iteratld atotg part of length (Fig' 71b)' 40

'\\ I [', t/

\"/ Y

Fig. 7I. Dorsal view of carapaces. a, }reoneetes hobbsi.; b, 0, P' palmerii c, 0. eotwressusi d, 0. alabanensis.

88 40(39) Central projection of first pleopod comprising more than 1/3 total length of appendage (Fig. 70b; see also Fig. 71b): )reonectes palmeri, paLmert (Faxon, 1884) (Streans in the lower Mississippi ValLey in western Tennessee, Mississippi, and Louisiana, and in eastern Missouri and Arkansas. Literature: Penn, 1957) Central projection of first pleopod comprising less than 1/3 total length of appendage (Fig. 70c,d) ...... 41

41(40) First pleopod with weak shoulder on cephalic surface at base of central projection (Fig. 70c): 2reonectes paLmert ereolanus (Creaser, 1933) (Streans in the Lake Pontchartrain watershed and the Pearl and Pascagoul.a river systens in Louisiana and Mississippi. Literature: Penn, 1957) First pleopod with no trace of shoulder on cephalic surface at base of central projection (Fig. 70d): 1reoneetes paLmeri Longimmtus (Faxon, 1898) (Western strean tributaries of the lr{ississippi River frorn the Arkansas River to the Gulf of Mexico, and streans westward to the Guadelupe River in Texas. Literature: Penn, 1957) 42(38) Rostrun with nedian carina; areola less than 5 times longer than broad (Fig. 71c,d) ...... 43 Rostrwn Lacking nedian carina; areola more than 5 tines longer than broad...... 44 43(42) Body strongly cornpressed laterally (Fig. 7Lc); distal por- tion of central projection of first pleopod not strongly recurved (Fig. 70e): 1reoneetes eorrpressus (Faxon, 1884) (Streans in the Tennessee, Cumberland, and Barren river systens in Alabana, Kentucky, Mississippi, and Tennessee. Literature: Rhoades, L944a) Body not strongly compressed laterally (Fig. 71d); distal portion of central projection of first pleopod strongly recurved (Fig. 70f): 2reoneetes aLabanensis (Faxon, 1884) (Streans in the Tennessee River systen in the vicinity of the Alabana-Mississippi-Tennessee border. Literature: Faxon, f885)

44(42) Mesial process of first pleopod with one or more proninences sLightly proxinal to caudodistal extremity (Fig. 70g): 2reonectes qtradruneus (Creaser, 1933) (Strearns in the headwaters of the St. Francis River in Iron, St. Genevieve, and Madison counties, Missouri. Literature: Creaser, I934a; Wil.lians, 1954a) Mesial. process of first pLeopod with no proninences slightly proxinal to caudodistal extrenity (Fig. 72).. 45 Fig. 72. Mesial view of left first pLeopods. a, 0reoneetes rTtpad.esi! b, 0. oalidus; c, 0. nais; d, 0' m'. meekii e, 0. m. breuisi f , 0. Longidi-gitus; E, 0. punctimmt'us; h, 0. oi,rtLis. 45(44) Distal L/4 of mesial process of first pleopod abruptly recurved caudally (Fig. 72;,b) """ 46 Distal L/4 of mesial Process of first pleopod not abruptly re- curved caudally (fig. 72c-h) "" 47 46(45) Principal. axis of first pleopod almost straight (Fig. !?"), }reoneetes nhoadeszl Hobbs, 1949 (Strearn tributaries of the Cunberland, Duck, and Tennessee rivers in Tennessee. Literature: Hobbs, 1949) Principal axis of first pleopod inclined caudally (Fig. 72b): Oreoneetes ualidus (Faxon, 1914) (Strean tributaries of the Tennessee River in ALabana and southern Tennessee. Literature: Faxon, 1914) 47(45) CentraL projection of first pleopod conprising less than L/3 totaL length of appendage (Fig. 72c,e) (For nethod of neasuring, see Fig. 3c).. 48 Central proiection oi first pleopod comprising nore than L/3 total length of appendage (Fig. 72d, f-h) 49

90 48(47) Chel.a _with conspicuous tuft of setae at base of fixed finger and with subserrate rows of tubercles on nesial surfaces of paLn and dactyl (Fig. T3a; see atso Fig. l2c): 2reoneetes rlais (Faxon, 1885) (Strearns in and Texas eastward to Arkansas; linits of its range not clearly defined for apparently often confused with 0. uirtLis. Literature: Creaser and Ortenberger, 1933; Willians, 1954a; willians and Leonard, 19s2) chela without conspicuous tuft of setae at base of fixed finger and tubercles on nesial surfaces of paln and dactyl subsquanous (See also Fig. T2e)z )veoneetes meeki bretsis Willians, lgiz (Strearns in the upper Arkansas River drainage in eastern Oklahona and northwestern Arkansas. Literature: Willians, 1.954a)

49(47) Dactyl of chela approxinately 3 tines length of nesial margin of paln (Fig. 73c; see also Fig. T2f): }z,conectes Longi.digitus (Faxon, l8g8) (Tributaries of the White and Little Red rivers in Missouri and Arkansas. Literature: WiLLians, 1954a) DactyL of chela distinctl.y less than 3 tines length of nesial nargin of paln (Fig. 7Tb,d,e)... S0

50(49) central projection of first preopod conprising aLnost L/2 length of appendage (Fig. T2g; see al.so Fig. 73d)z 2reoneetes ptmctimartus (Creaser, 1933) (streams in the Missouri, st. Fiancis, and white rivers in Arkansas and Missouri. Literature: Wi1liams, 1954a) Central projection of first pleopod conprising nuch less than I/2 Length of appendage (Fig. T2d,h). ' 51(50) Mesial process of first pleopod rather strongLy recurved (Fig. 72d); areola constituting less than s6 per cent of total length of carapace; tubercles on nesial surfaces of palm and dactyl subsquanous (Fig. 73b): 1reoneetes neeki neeki (Faxon, fSg8) (Streans in the Arkansas, Red, and Wtrite river systens in Arkansas. Literature: Wi1lians, 1954a) Mesial process of first pleopod only noderately recurved (Fig. 72h); areoLa constituting nore than 36 per cent of total length of carapace; tubercles on mesiaL surfaces of paln and dactyl distincly eLevated (Fig. ZSe)z 2reoneetes oiriLis (Hagen, 1870) (Lentic and lotic habitats in Canada and the norihern part of the united states; the southern boundary of its range is not known. Its reLationships to O. nai.s arrd 0, ecuseyi., the Latter here considered a synonym, are far fron clear. rntroductions into Maryland, california, and perhaps else- where have resulted in its being one of the nost widety dispersed crayfishes in North Anerica. Literature: Crocker and Barr, 1968)

91 Cataloe of Crayfish Snecimens

Annendix C t[{au + tr Nortln F'L oe Eaetfork oP 6\ar,l-rlvc.r Cr.,5f;sln e.tt

*.1!t Fo.'.^ rrnates t t 2$Foc'n rnrat<*s al fs{r*i li nna,te S ?,8 F-,--t

S, b Fo"--.131 C-*.bacr Fra€- Go^vS '.Or c"n 4& 37<.<.;.-s, Virl\< + G.* I ld -*[<-<=.i]ttlil. ft AND Fon-.^ wro\e3:111 ' { ..VoY n.o.!"<= 3 ifotal nrrr'tz(': l S , Fa.^o,ks" 23 toY F-"^^IeS: 3 26 fofa.\ Fer ^^leS =

fota.-\ : t{ | nlroiab * 3 A2-, Coehig<- Cov.'t1 1.2 .\ t/-r yn'.. Sa,n ?--elro R''ver Ca. Jo,^r,,.rg!..rr,,zra FFo-v,-r Chc,'\esttn Bni J3<. 4,ts,eatio-.n 3

3g'c.e.\ c,5; . V;c;\ i S l*€o."..n ^a'l<- = I

F<-r-.a,lcs: I

-tfr"l

AZ, Gila,-P;na-[ Covntie-s Gila. R-i*re.r, fr6t bc-lou) rohFtuan<.z uTI\^: 5q1-rcoE 34LB t€o \,

ttfarlq3 R. tJ. C-larkssn '' rr\-' s vl 13 a t c'3 4 t n:}= S;ncc- r- r'a s Fo*:l1t 1 Ca^b a,r', Jae $-*Jo oJtr.'s VoY tr+a't+3, @r pos'*rrre- &qusa t,tr : Ca-rt-ae \ a, a 4 br' t^ S..ibF+*. s?3€-ica i.Gl . eswlJ no* ^"&'up;+l^o.,zF Gen.rs', Occ-onc&os 4 F.rtt $-r.a -^.ot<.Ct{.bbr lela), F{or.re^r-- srrs'g<.r-.f dtl bvr SPec.r'as: Vi |-:\i S .t tL.a*. c-ra,6 F;a\.co *rcO-re*: Orc-oncc:te3 =S ^s{ nn cFl'obto' l(r1)- \{o.^zve6 3Fvziat.. yi c.\.'S *& J. qr* Fo.rt1 Co-'FrJ+-t4 ru.* 3A B"^^ ana\q6: 2 bo+x*€Q.@vdiS'

1 e'l, i, satt\c.s rc$ a la-ora # 7 // jf ps+e: Q6.a- Iagc- ,!, n*f Foe.^. -no-[<- :,n Aa Cp-c,ha, C-o. Unn o..o*d te: br1--a -taf leP2. t,r,.,;th, 4rrlL+ l$ C,{,te.-lo. f1..;3 v l/, tr.lc.s* srf *\r'

Ag Fsr.^ *a,te::S fotr-l= C tlnlqd * t2- / AZ, APcchc- Co-'*3, u..lest FsrF oF tsta<-{

J@ Co.n^ v,rra,tc.: I

<-''n*a? ,/ ^rlrr ll6 it t3 'l'2' -/ -,lul

Fa,m: I 14 : C-^-h'a"rivrc'r- 3..1b \/ 6c,vtvS'. O c eo^t'c'crk6 9e<<;e'3i r.J) Iti\r 3

19& fo"". r'^Jc-= | nlnllb ''t{o / AZ. t-a. Paz- C..o,*\ C;u,.,la i'll^JR - fftif' L.atoez- F"^J (q) A crlFrs\ 5-as-a6 + 6-aq-14

Q,olLoczfor.rt I Lo.Bo'.bqn'- 11\sr5 sn 1

Fa-^i19t : Ca*b,,..i Aa'e J 6ubfa'^ I \1 : C^-l6ar',6a€- J Gaqrs3: Proc^.nba,rtX

Sub' Ge."t vS I S<,^t",v[i caxr br.r.vs

5rp-- \<"s L C-la' k-l i

l* n'rol.-s= 1 r..e-inir,1 J;lr \,&tw-{,6 jw*^;1a- qarL< J larb F'* ,*.rLr * (1 :rvsb s*^^+e J r f r rf tc, / Az-. m*-iq.oga. C.o. Ca*P C7e'e,'lr.-_ 3'aS-' Fco- J\ ain rv.A lvlea C*tle<:te-zl bg . l- a.r*ran , P. 5!oarKa L-^; \.4'. Co- 6o.").e- J In Ca"- b^"^o €- S.rbFa. ^: { ; 6 e-w') S j Or-co-.<. .t(-s

3 j+etit-6 I \7: ": [;s llt 5r^m ales- . lA S.-. tl.ol.es t Z alcs = I Suva^:\<- 'aT fc*^[ rna,a,[<-3 ' 9 Fo*-'l.z= I ls5'l- 6 '\la

f...^ -lra" 7 11:o/1a # t1 fra. 4eea.nl<^c- C,o. Bcra,r,rc- C-re(rL Fc

Co ttc"z;tors'. B. Beg\+J' R. r^':. C-tq rLso vr' D. For,*er, 6, ScJ^.fS vnni l\cr

Fa.*. lJ,Co'-rbari J a c- S-b F--; lf I *{o aF r Ae L .l Ca 3. r^ u s'. Oi* -,e-c,Fcs, Spqsle3'. Vi crlir l$8r".^ rnnorles t 2 JCD Gort'. *vr"lc<= $ Tc{-r,t- J 1l:i. _,. lq6 #:o 20 FC.vn 1,8,1 Az. G-

JSSqc.,1 r.nr.t.( " 2 Sg.f ur:-\

Fa.,-.111'. C.e.,^^hqr i Je... \) ?le Uop : s;ne.c- us Flt ts.".. ,r,...,J<.< rrrgr.<- SrbFa.*ni \1 trra c- \r' Ca.^'-l5rr; Pt\r €,F.* l-r {\,ra Sa*pl .{ a G<.--r-5'. Q g c,o,"".. cltcs Pos;\ive- S D c,cwl of ,^.."+ ta r.,o"l.-1. 3V<-;c-s Vir.i\i< hto...,*revec, oJ). q^! p{r$q$h 40 F*. ,*.,fcs. 5 Orc^-a-c*r^r V-.gilS- F.,,.^..Jas. 3 ae1*t: f,

Slalqa +e2 9e*l na3iJ dar.' l<.f| gl-rp..l { e-:' 5^.L l$ Gc.",.-.t<, Az.. Ya.l*g4l--e-o. 4r:r'+;*e ? n

Far..r,r.; l,{ j Ca*boc\de,e J 3rbF-.^l 11 i 1a**c^r'v\1e J 6*" 11 r..r11', O. *^ -+:- -;:ge.ei cS . gi cr\.9

ls F,^-^ -*le{: 2 F... ^'-t<-s = 2

\.; \_:j ! rr n, , It ,ti' ':\? .__-_4 / l t i t. \

5Ll fte^ r),^Ff tlE.-- "^"1 : 3+w:w55 s(qlqb + 13 trf\ AZ-. YV1a"i<-$Fc , -'\{0o :t*qo 1'(e €^.\.^ C-o.-o-t betr^rae,wr Al Me. l; st--i na'*- J 6<4s5. ?ro c-a*\rae rzS s ub 6c-r,rvs . S <-^pv\; -^,.,^b.lys Sgac.,a<,', C-\-rrk-:i I* ,r,.a,\as . O

Fc.*ale s : I

tltzlnu fiaL{ Az-. J\Aar'.a?a C-o,rnt1 . f*.^a4<- C,-.lU--^ C^-.-[ bo{*.o'.. A-A\\lr|ar$Rur^t q-et. on L/nnl,,te".s; \ c-r,tl 1<*<-A lo-.,\ , f. lfvrvna.r oc*al: tlqlqb \J Fo*.'f . Fo-:13: J C-^ ".b a,rtJac- S,rLs Far-; ll: C-.r^^h<^;"rL€- Ge--tus'.

Go.n,ua I ftoca-.^.bar,S . Sp-a-c-; t5'.

v< : 1vV Gnn S- a tuli 6,-n1l6lr'4 9 p-e;c-s'. f-tcrki i

I 5j rrvf o.[e-< t I 1Ft Jr{9 6natas : 3 ?rn-nl', flttl

A1;J.engcJces U, .t !:: ,

- D f. Ar--zra..rn TL.'80.,-. t 7s-*. * 2-l

AZ . G.=-^lcr.- C,o. T/R- T \.oS R aE. o S 3e<_., 32 S p<-a; F;<- s,,*r q Ls<.a*;on. €-5[.- Qeeel- €- 6^S,-3 s*-'tion &rlaebr-.J Ae,f D Uo-So. 6-^.1^ \: ^- D.a'tc." lqouoc- # Nt"fa .aT,4aL 44._ A pcobta^-f,.€_ F,e-LJ*: MALoo.qlo66 t-*g o' c6*$F,sl^.a-a, 1-ho: r v4e-4svr*^on'44 na;*\st (-a*o\ u1u-d b.. : A . Msorhovs.e- F;{' llotbs !.!^{-ase1<-r14-..da frt- J Ao A u to C' PL r'a-*r o', r d,s J{4- Ft+ Lr^ 3 ers F**; 15 . C-^b ar, Je-€ fiee'!,ral\,e6*5For O, U,.tt,: , BOfq 5',,!r F-'.,+l 13 : na"e- Ca -l5eri S€^s otr €-3t<. Cc-.-t- C*UF,'C\ ;1er Ge,yr r.rs : Qccrn<-c*<-S; t*-" \i k-a- tl^iS. 3gz{te'3: tJ: r; ti s l$ 8",^^ vnAatcs'< { A€Ec*,.r.-tett l {t'-^te.g, = 5

*rs AZ. Va'ogo: cs. Verd,z, R;var aG's-c- hpcr-Shoe-. Rc,o. gatt

f *Fo"rr, ,n ol<-= I eXs*nT?- JE R"- *rle. = I F-.'-o,\.. I #24

AZ- rrrfsL.o.r<- CO. -T/R.:llN l'?W Se<: S9 =;'c.c-r,9i<-'.S€4 [Je'{

,oea+nyr: 8;ll llr.rill;awrs R. ao^pl, o.r;th f\;vlgeal WaSl, &,lt+el-A bl: Aato LJo-tg-'ts- R'r*'"te'A q4 o tos D"+s *.N# J\is is q {-ext+fir,e a.e, sin

Fo*o"la= I

t3p Az. Yauap-l 3r. r/e.:r\'l.oN R3.o € sc-<.i'? spc<.tf .e', S€ rt Sg3'l c-a' Lq<.o,t icn : V e-r A + Rirre r Q €5 c'a'ra^'r€ h !'Ya t'Jor^5a'^r'<- A69D Beo^of' T c ot\<-42-J b3: #}}*.-. Song C{l^r. ^\eosur+}tzliahPl : vt4*el^ou9z- Q,etato3rr-c-,{ \ A. *rr.t*c.n $csnn+\4- I $ f**.^,r,ol<' *t F^*rt b.-'^\*rrA^- ur+- b.- c.*^^d-{e,t1 c.cura*e du<' Suh€a-,lX'. C*qbar i aa€- .}5 -+\- Fns <..q,rJ;rYqr oF+\& (qccrurc6. 6o^-5 1 Qrc--.. q-c.,fc3

Spe,c-;as ', ! i r' i \; S tisr-' n^atc= \ A@R..^ ynate < I

Lr,.al< = I

F3l Az.Pi,,r.r*eo. -lF/R. t3S lSg Sac-:14-3 Loc.ac+rsv11 Sa-!;-o 3',<_<*-r bu i A . " Dartt:9cttstB ^-.i1|l;1'il*- # Itl"t-'. -{l'.is is onll < *-^}J'v<-alD F'4d *: F<.osso D,r<-{s +f^r- Foor a-q61.{1'trn oFt\\a fa,^:l &*5ar; J1: 6/"rc- 3 m.1A 5s,zrr oF Tk Pe.ei -J \,cr,r- SvhF-; 11 : (--.-,ban,naC yta<-43sr!-c"c.nhs lO_<-,nccc.^rmrbg, J ^- U G e.r:, vs : O Fc-o"r.

l-g5a,'4;.e.1i N-!c:osg (,F

Fo^ ; I .1'. C^*to *' C, oe- U 3wb Fa^ llq : C-o *b^r i r\e.€. $ G e- wus'. O r6,.rnc.eta3 S?ra<,l<-S', U;'-:l i6 o?]9! 9ctrr/. ,nna-la3:3

*3.{ Az. Va..l-pq', co, r/c', t5.oN Ra.oE S<.cr3r.{ sp<.<.1ti<.'.N€-q N€.{ Lor-*rqn:Watkcr Crc..K, v1€-ar \-J

lliF.c"^ *a\es= I &*& Fsc"n vrna\ef r {

Jii,rvr-^ll. rrle,\+ r I

Fa-,.*te: I

Srfr.lov*, IC- Fr.rn^^lc-. I

#30 AZ. Cocrrn.ns C-o' T/F-.-ft9.o N Q.. U.Og sec,,', a4 sp<-e.9;c-r NEtl €Eel l-ee{;cn'. fa*t C-t<-or ejce.Ll F\sns.r_ C-..-sst ig c,or\e.cbe-at bt i Aogo NJ.^gc.^e bn--.\ D4+<.d i qqoba,q F,JJ * ', bBDeolqq 06"''l 43tgLolv<'4'trl| ' rr I. r\, V\p'srhwse- F.,'^^:ll: Cr*"- '' Jo? F-.*l\r4'. !\^ a 3 $vL { Co.-loar 6or^v: I Or<'qtrJas Spc<-'c4; glrril;s f 2BR.**clq. . | @- loJrna'nr Fo',.^*l.s' Z @ lo-l vnv-r {; {4r""r'n tt3'1 SqJtl Az, Co.-o-'.no Co, r/*"f lq.O N F.ll'og sc'ci 35 sf?<,.F,'c,: Swt-{ q Loc.alion ; G.a.E\ C)*r Cee

3pc.<-reS'. V;c:\5 [* Fo.*. natc cL @ S9*-+c,^75,-.4

Sc,r^^,tcs . L Q- b4't*^t 33'^'^ tsg

,52 . Vrvapoi Co. r/e; r rf . orJ R.t.o g Sq<..! J? SPz<-.4;<-" Nk,4 U€&{ Loc--t'sn '. *<,ll Pst^l c o'.$\eea c(- c40-Lcka bg.. Ac'nD WOrbA^JlDgD T[.,]: 'E^tcoA^{'v<- f D Ddr€l : q3to13 @. F'rs{ Fo"-t ---tsg u'1,-r<- F,.la+ : bgboq3l3lot3 J:rn4.-z-,"t tJ\ b''!JJYa;1 ' &lolo5vJ b1: B. De,nqur 9^nt-'^"tt-:- F-.^, t3 : C-.-ba c', Ja.a s;,,,rbF...,-^,i13'. C-*-^\ ar , rna{-

C=<--r v5 i CfE,ott..,{cs Spec-;4s I Uiril;s &NEFe.-r. ana.atct I @c{t'^""^ F-.-'..o,\-'l @b9,^a R3{ 41. Vav.g^. Co' 'r/n:rtr'oN R-t'od 3*'<"r3? Spc.c.i$rc- i,r.J/A .5n;l'<-lo.-h*'' lls ctgr c'rosstlS j7ljg)kei -il\,i3 g"o1F I sh L,- s s*cm$'<- Lo.-tio-z ,ft^, 3Y +L& tk o *..+al f ontr t^ c-ot\ e4e2 lo^,. : AcFO tsPDrOAw't bSO -t :. .FNvm+

5 Pe,cics '. Vtcilt s l*Fo"-6.t-l e: le- q'6 rmtm +43 A"-. n a.r; ceP Q.o. B.-"d.slv-1 C^n*l /u -,. no-L\ og g-q-Sce<- H'^,u1 ^ J u c''L\a-c,.a+e.-A . t/zY lqr ( ,r to-,-.' l)','^ C .. 6\-bo,. ")qz I vlo F^-.' lq I Co-r.L4r..rnq.e J 6z.rzrv 3 I GC,-4 e.-1eS SFc.c,t.-'!ini\.t $ ; 9..-, ....av\e . I r^atte a\b 9o. r,r = F.,-^-te-.3 *qt gr(. <,k *€tt : ot ,4o.2. W\6-i<-sFq Cs' P:*<-o lv\c..Cli^$, e-'nc-ak4 tA{ {' 5n'w--r^ J i 5 a *'""*'*t'"'e' D&.-4 : tlellb * Uot<"' @t-t{r-''gh'tl^1 3 aS 4 Sitsh Pbt* nt-l-<- F-*' lq : C-a*r'tat ia\a'c- ..{e-^t"Fic-t io\^ J b-4'- v\qr. aval\et'l i t Sac'n, .3 1"t '' UJa.s -, 5 -bF* ;l 1 C.^-..b af rt *e 5 e ?' s-$5 n. s{ o 9. P=sr,^d351: G*v...-,5 I P-oqo*tr11 -S a3 il \^ra-s aJva,--J';'J'o{ +o to-, oF k-''i t\'<-di3*riLv'te' \^r<"s o*'t I C-(4" Posg,b-U P<.-<-i<-3 " o'^l' s-'*f' ts Pe=.=*]=ns c--*lU' 2t'LQ' 6n".lc- -- I .L

4v4:ta-L(c.' o-{- +\^. pc-'b *tsre"' The Known "Wild" Crayfishes of Arizona:

Distribution map

Annendix D The Crayfishes of Arizona The Cravfishes Known to Neighbor and Border Arizona:

Species list and distribution Map

Annendix E The tr(nown Craylishes of Arizonq Califomiq Coloradq , New Mexicq Utah, and Mexico

* (I)=hboduced (N)=Native ?:Suryecedintroducim

Arizom Medco (l) Orconectes irilis (N)Cambarellus alvarezi (l) Procambarus clarkii (N)Cambarellus areolatus ( N ) Cambarellus chapalanus C.alifomh (N ) Cambarellus chihuahuae (N\Pacifastacas fortis (N)Cambarellus montezumae {l)2 Pacifasncas gambehi (N)Cambarellus occidentalis {N ) Pacifastacas leniusculus klamathensis (N)Cambarellus paacuafen$s (l) Pacifastacas Leniuscabs leniuscalus (N)Cambarelfus prolints (l) Pacifastacus leniuscalus tlowbridgii (N)Cambarellus zempoalensis (N )Pacifasacus nigrcscens (N)Procambarus acanthophonts (l)Orconectes virilis (N ) Procambarus bouiei (l)Procambarus aa&rs (N ) Procambarus caballeroi (l\Procambarut clarHi (N)Procambarus clarhii (N) Procambarus contrelasi

Colordo (N ) Procambarus digaeti (N)Cambarus diogenes (N) Procambarus elichsori {N) Orconectes causeyi (N ) Procambarus hofmanni (N) Orconecbs imfiunis (N ) Pracambarus hotonhobbsi (N)Orconectes neglecfus negbctus (N ) Procambarus llamasi (N)Orconectes vililis (N)Procambarus fi6icafi8

(N)Procambaras simulans (N ) Procambarus mirandai (N ) Procambarus ortmannii Neryede (N) Procambarus paradonts

(N)Paciftstacas gambelii (N ) Procambarus pilasimarus (l)Pacifasacas leniusc'ttlus leniu scalu s (N) Procambarus riojai

(l) Paerftstacas leniascrlus trowbidgi i (N ) Procambarus rodiguezi (l) Procambarus clarhii (N ) Procambarus ruthveri (N ) Procambaras ruthteni zapoarynsis NenMedco (N ) Procambaras s i mulan s re gi omont a nu s (N)Orconectes deanae (N)Procambarus tsiutlanentis

(l) Orconectes rusticas (N ) Procambarus tlapacoyanensis (l)? Orconectes virilis (N ) Procambarus toltecae (l) Procambarus clzrbii (N ) Procambaras 9arzquezaze (N)Procambatzs simulans (N)Procambarus veracruzamts

(N ) Procambarus zihuateutknsis Utah (N ) Procambarus sbordonii (N)Pacifastaats gambelii (N ) Procambarus acuhts caevachicae (l)I Pacifastacas k ni a s cttla s leniu s calu s (N ) Procambarus gonopodocristatus (N ) Orconectes virilis (N ) Procambarus villalobosi (l) Procambants clarkii (N) Procambarus nlitlae {N ) Procambarus roberti (N ) Procambaras stenthi (N)ProcambaTus cueEalanae

(N ) Procambarus xpchitknae (N)Procambarus oaracae oaxacae

(N ) Procambants ouncae reddelli

Taylor, C.A; et al. 1996. Cqsvatim statrn of Crayfishe of tbe Unir€d Stat$ erd Csuda Fishtri€s Vol. 2l (4). Aneriru Fbhsis Sciety. Beth6d4 MD.

Hobbr, IlH. Ir. 19?2. Biota of Frc*mataEcsystm. Idanifi€ti@ Mmul No. 9.; @he (Astmidm) ofNcth md Mddle Amqie. Envimmomal Protectim Agmcy Proj@t # I 8050 ELD, Cootra* # I +l 2494 U.S. Govmant Printing OEc. Washingtoa D.C.

Eobbs, H. E. Jr. 1989. An illwffied ChecHist ofthc Amim Crayfishes ( D*apoda: Astaci&e, Cmbcidae mdPffistacide) Smitlsmim Cqdrihrti@ to Z@log. Nwbu 480. Smithmim Institutim Prw. Wartrington, D.C. The Crayfisbes Known to Border Arizona