Ambrysus californicus Montandon

(FIGs. 2–3)

Ambrysus californicus Montandon 1897: Verh. Zool.-Bot. Ges. Wien, 47: 12, 18–19 (original description, key). A. californicus: Torre-Bueno 1906, Proc. Entomol. Soc. Wash., 8: 54 (list of taxa).

A. californicus: Kirkaldy & Torre-Bueno 1909, Proc. Entomol. Soc. Wash., 10: 186 (list of taxa).

A. californicus: Montandon 1909, Bull. Soc. Sci. Buc.-Roum., 17: 320 (key).

A. californicus: Van Duzee 1916, Check List Hem., p. 52 (list of taxa).

A. californicus: Van Duzee 1917, Univ. Calif. Publ. Entom., 2: 458 (list of taxa).

A. californicus: Hungerford 1920, Univ. Kans. Sci. Bull.,11: 199, 204 (key, translation of original description).

A. bohartorum Usinger 1946: Univ. Kans. Sci. Bull., 31: 195–196 (original description).

A. californicus: Usinger 1946, Univ. Kans. Sci. Bull., 31: 196 (morphology).

A. bohartorum: La Rivers 1949, Bull. So. Calif. Acad. Sci., 47: 108 (key).

A. californicus: La Rivers 1949, Bull. So. Calif. Acad. Sci., 47: 108 (key).

A. californicus: De Carlo 1950, An. Soc. Cient. Argent., 150: 7 (list of taxa, distribution).

A. c. bohartorum: La Rivers 1951, Univ. Calif. Publ. Entomol., 8: 309–310, 336 (subspecies status, habitus photo) (NEW SYNONYM).

A. c. californicus: La Rivers 1951, Univ. Calif. Publ. Entomol., 8: 306–309, 336 (supplemental description, habitus photo).

A. californicus: Usinger 1956, Aquat. Insects Calif., pp. 201 (morphology).

A. c. bohartorum: Usinger 1956, Aquat. Insects Calif., pp. 202–203 (key).

A. c. californicus: Usinger 1956, Aquat. Insects Calif., pp. 202–203 (key).

A. californicus: La Rivers 1965, Biol. Soc. Nev. Occas. Pap., 4: 5 (subgeneric assignment, distribution).

A. c. bohartorum: La Rivers 1971, Biol. Soc. Nev. Mem., 2: 66 (list of taxa).

A. c. californicus: La Rivers 1971, Biol. Soc. Nev. Mem., 2: 66 (list of taxa).

A. c. bohartorum: Polhemus 1979, Calif. Insect Surv. Bull., 21: 134–136 (key, distribution).

A. c. californicus: Polhemus 1979, Calif. Insect Surv. Bull., 21: 134–136 (key, distribution).

A. californicus: Polhemus & Polhemus 1988, Catal., p. 523 (list of taxa).

A. c. bohartorum: Polhemus & Polhemus 1988, Catal., p. 523 (list of taxa).

A. c. californicus: Polhemus & Polhemus 1988, Catal., p. 523 (list of taxa).

A. californicus: Dinger et al. 2005, Southwest. Nat., 50: 242 (list of taxa).

Diagnosis. ThIs spEcIEs cAN bE dIsTINGuIshEd fROm OThERs IN ThE cOmpLEx by ThE SGP pOsTERIOR mARGIN wITh A fOuR- LObEd AppEARANcE, wITh TwO cENTRAL LObEs ANd ROuNdEd pOsTEROLATERAL cORNERs. ThE cENTRAL LObEs ARE sLIGhTLy pROducEd pOsTERIORLy fuRThER ThAN ThE pOsTEROLATERAL cORNERs (FIG. 3F). ThE LATERAL mARGINs Of ThE SGP ARE NOT REfLExEd ANd ThE pOsTERIOR mARGIN Of LATEROsTERNITE VI Is sTRONGLy cONvEx (FIG. 3F). ThE mALE AGP Is EvENLy cuRvEd TO ThE RIGhT fROm bAsE TO ApEx ANd wIdEsT IN ThE bAsAL 2/3 (FIG. 3B). ThE psEudOpARAmEREs hAvE A mEdIAL cONvExITy ANd dO NOT TApER TO AN ApIcAL LObE; ThE pOsTEROmEsAL cORNERs Of ThE psEudOpARAmEREs ARE ROuNdEd (FIG. 3C). ThE scuTELLum hAs A dIsTINcT dARK bROwN dENdRITIc pATTERN ON A mEdIum bROwN bAcKGROuNd (FIG. 3A).

Discussion. ThIs spEcIEs wAs dEscRIbEd (MONTANdON 1897) bAsEd ON A sINGLE mALE spEcImEN cOLLEcTEd IN sOuThERN CALIfORNIA (UNITEd STATEs). IN 1946, UsINGER dEscRIbEd A. bohartorum fROm SONOmA COuNTy IN NORThERN CALIfORNIA (UNITEd STATEs) ANd suGGEsTEd ThAT ThE spEcIEs wAs sImILAR TO A. californicus, buT dIffEREd IN hAvING A bROAdER pRONOTum, wIdER EmbOLIA, ANd AcuTE pOsTEROLATERAL cORNERs Of AbdOmINAL TERGA III –V. HE cONsIdEREd ThE AcuTE pOsTEROLATERAL cORNERs TO bE sImILAR IN AppEARANcE TO ThOsE Of A. mormon MONTANdON. LA RIvERs (1951) ExAmINEd spEcImENs Of A. bohartorum ANd fOuNd ThAT ThE mORphOLOGIEs Of ThE mALE AGP ANd fEmALE SGP wERE IdENTIcAL TO ThAT Of A. californicus; ThEREfORE, hE pROpOsEd A. bohartorum TO bE A subspEcIEs Of A. californicus, wITh ThE AcuTE pOsTEROLATERAL AbdOmINAL cORNERs As ThE dIAGNOsTIc fEATuRE fOR A. californicus bohartorum UsINGER. AmONG spEcIEs Of Ambrysus, vARIOus ExTERNAL mORphOLOGIcAL chARAcTERs dIffER IN sIzE AmONG cONspEcIfIc pOpuLATIONs ANd hAvE bEEN usEd TO dETERmINE subspEcIEs bOuNdARIEs. SITEs ANd WILLIG (2000) ANALyzEd ExTERNAL mENsuRAL fEATuREs AmONG 13 pOpuLATIONs Of A. mormon ANd fOuNd ThAT ThE REsuLTs dId NOT suppORT ThE EsTAbLIshEd subspEcIfIc dEsIGNATIONs. ThE vARIATION IN LENGTh Of ThE pOsTEROLATERAL cORNERs Of ThE AbdOmINAL sEGmENTs IN spEcImENs Of A. californicus REpREsENTs TypIcAL vARIATION, Of whAT wE cONsIdER A NON-dIAGNOsTIc fEATuRE, ThROuGhOuT ThE dIsTRIbuTION Of ThE spEcIEs. ALsO, cONsIdERING ThE subspEcIEs As A GEOGRAphIcAL vARIETy (MAyR 1982) wITh mORphOLOGIcAL uNIfORmITy IN ONE OR A fEw dIAGNOsTIc chARAcTERs (WILsON & BROwN 1953), ThE subspEcIEs Of A. californicus shOuLd bE IdENTIfIAbLE ANd AssOcIATEd TO A pARTIcuLAR GEOGRAphIc AREA IN CALIfORNIA bAsEd ON ThE LENGTh Of ThE pOsTEROLATERAL cORNERs Of ThE AbdOmINAL sEGmENTs ANd ThERE shOuLd NOT bE OvERLAppING Of ThE RANGEs. ThIs Is NOT ThE cAsE bEcAusE ThE spEcIEs hAs A cONTINuOus dIsTRIbuTION IN wEsTERN CALIfORNIA (sEE ALsO dIsTRIbuTION mAp IN POLhEmus 1979) ANd pOpuLATIONs wITh bOTh cONdITIONs Of ThE pOsTEROLATERAL cORNERs Of ThE AbdOmINAL sEGmENTs OccuR (E.G., SAN LuIs ObIspO CO.). Thus, bEcAusE bOTh chARAcTER sTATEs ARE pREsENT wIThIN pOpuLATIONs, wE cONsIdER ThE dEsIGNATION Of ThE subspEcIEs NOT vALId ANd pROpOsE ThE syNONymy Of A. californicus bohartorum wITh ThE NOmINATE subspEcIEs.

Variation. NO vARIATION wAs ObsERvEd IN ThE dIAGNOsTIc fEATuREs.

Type locality. UNITEd STATEs: CALIfORNIA.

Repository. HOLOTypE Is hOusEd AT NHRS.

Distribution. ThIs spEcIEs Is ENdEmIc TO ThE wEsTERN pART Of ThE sTATE Of CALIfORNIA (UNITEd STATEs), whIch Is IN ThE PAcIfIc BORdER physIOGRAphIc pROvINcE (FIG. 2). DINGER et al. (2005) REpORTEd ThIs spEcIEs fROm 10 LOcALITIEs IN ThE CuATRO CIéNEGAs bAsIN IN ThE NORThERN MExIcAN sTATE Of COAhuILA. ERIc C. DINGER KINdLy ALLOwEd us TO ExAmINE sOmE Of ThE spEcImENs Of NAucORIdAE cOLLEcTEd IN TwO LOcALITIEs (PuENTE ChIquITA ANd RíO GARAbATAL) IN ThE CuATRO CIéNEGAs bAsIN. WE fOuNd ThAT ThOsE spEcImENs ARE NOT Ambrysus californicus buT INsTEAd ARE Ambrysus lunatus

UsINGER ANd A. pulchellus. WE AssERT ThAT A. californicus Is NOT dIsTRIbuTEd IN MExIcO; ThIs spEcIEs Is REsTRIcTEd TO ThE cOAsTAL RANGEs Of ThE wEsTERN sTATE Of CALIfORNIA IN ThE UNITEd STATEs.

Published records. UNITEd STATEs: CALIfORNIA; MéxIcO: COAhuILA (MONTANdON 1897; LA RIvERs 1951, 1971; POLhEmus 1979; DINGER et al. 2005).

Type material examined. HOLOTYPE ♂ (by mONOTypy): [UNITED STATES: California]: S. CALIfORNIA / MORRIsON / Typus / Ambrysus californicus MONTANd. TypE. HOLOTYPE ♀ (by ORIGINAL dEsIGNATION): [UNITED STATES: California]: SONOmA CO., CALIf., AusTIN CREEK NR. CAzAdERO, IV-30 -[19]35 / R.M. & G.E. BOhART COLLs. / HOLOTYPE Ambrysus bohartorum UsINGER / CALIfORNIA AcAdEmy Of ScIENcEs TypE NO. 7159 (CAS).

Material examined. UNITED STATES: California: ALAmEdA CO., LIvERmORE, CALIf., X-1-56 / J.A. ChEmsAK cOLLEcTOR / Ambrysus californicus bohartorum USING. DET. MENKE / C.J. DRAKE ACCESSION (1Ƌ USNM); ALAMEDA CO., 17 MI SE LIVERMORE, ARROYO MOCHO, III-24-1976, FRED G. ANDREwS / ASSOCIATION MOSS IN STREAM (1♀ CSCA); COLusA CO. INdIAN CREEK 1.5 mI ON ROAd TO COOKs SpRINGs / SW Of LOdOGA 7-IV-1971, HuGh B. LEEch / Ambrysus californicus bohartorum USING. DET. J.T. POLHEMUS / C.J. DRAKE ACCESSION (1♀ USNM); COLUSA CO., MENDOCINO NAT. FOR., MILL CREEK CmpGd., 20 IV 2015, 495 m, MILL CREEK, N39°21.33' W122°39.32' (WDS-A-1981A) / ChERyL BARR AND WILLIAM D. SHEPARD (2Ƌ, 9♀ UMC); COLUSA CO., MENDOCINO NAT. FOR. MILL CREEK CMPGD. 20 IV 2015, 495 m, MILL CREEK, N39°21.33' W122°39.32' (WDS-A-1981B) / ChERyL BARR ANd WILLIAm D. ShEpARd (1♂, 3♀ UMC); COLusA CO., MENdOcINO NAT. FOR. REd BRIdGE, 20 IV 2015, 449 m, MdL. FK. STONEy CREEK, N39°22.64' W122°39.14' (WDS-A-1982) / WILLIAM D. SHEPARD, LEG. (1Ƌ UMC); COLUSA CO., MILL CREEK CPG., IV-30-1980, FREd G. ANdREws, swEEpING Quercus SP. [COLLECTING HABITAT MUST BE INCORRECT] (2Ƌ, 1♀ CSCA); CONTRA COSTA CO., CAL., AUG. 25 1911 / MARSH CREEK / BRIDwELL COLLECTION (1♀ USNM); SAME BUT / J.C. BRIDwELL (1♀ USNM); SAME buT / Ambrysus californicus MONTANDON 1897 (IRA LA RIVERS) (1Ƌ, 1♀ USNM); FRESNO CO., FIRST wATER IN dEscENdING LOs GATOs cREEK, 1.7 mI. SE Of dIvIdE (=SAN BENITO-FREscO CO. LINE) / TRIbuTARy Of SAN JOAquIN VALLEy, 6 OCTOBER 1941, W.I. FOLLET, F60-41, (1Ƌ, 2♀ UMC); FRESNO CO., WALTHAM CREEK BY ROUTE 198, 5.5 ROAD MILES SW. OF COALINGA / 27-III-1970, HUGH B. LEECH / C.J. DRAKE ACCESSION (1Ƌ, 1♀ USNM); SAME BUT / Ambrysus californicus bohartorum US. DET. J.T. POLHEMUS / C.J. DRAKE ACCESSION (1♀ USNM); LAKE CO., BEAR CR., CAL., IV- 13-1968 / A.J. GILBERT (1♀ CSCA); LAKE CO., N. FRK. CACHE CR. [ON] HwY. 20, CAL., IV-21-1958 / C.J. DRAKE ACCESSION (1Ƌ USNM); LAKE CO., STREAM FROM COMPLExION CANYON / BARTLETT SPRINGS, 6-IV-1971, HUGH B. LEECH / Ambrysus californicus bohartorum USING. DET. J.T. POLHEMUS / C.J. DRAKE ACCESSION (1Ƌ USNM); LAKE CO., 8 MI E CLEARLAKE OAKS, CACHE CRK., VII-3-1975, FRED G. ANDREwS (1Ƌ CSCA); LOS ANGELES CO., CAL. / COLLECTION COqUILLETT (1♀ USNM); SAME BUT / FROM COLL. USNM / 0 53 / Ambrysus californicus dET. MONTANdON 1909 (1♀ USNM); LOs ANGELEs CO., CAL. / COLLEcTION COquILLETT / 0 53 / Ambrysus californicus MONTANdON 1897 (IRA LA RIvERs) / Ambrysus californicus MONTD. DET. MONTANDON (1♀ USNM); L[OS] A[NGELES] CO., SAN GABRIEL CANYON, D. TIEmANN COLL. / U970 / L.J. MuchmORE, JAN. 26 1935 / C.J. DRAKE AccEssION / Ambrysus californicus MONTANDON 1897 (IRA LA RIVERS) (1♀ USNM); MENDOCI[NO] CALIF. / P.R. UHLER COLLECTION / Ambrysus californicus MONTANDON 1897 (IRA LA RIVERS) (1Ƌ USNM); SAME BUT / Ambrysus californicus MONTANdON 1897 dETERmINEd by IRA LA RIVERS '50 (1♀ USNM); MENDOCINO CO., NAVARRO CR., CALIF., VII-2-57, J. POwELL COLL. (1♀ USNM); MENdOcINO CO., NAvARRO RIvER, HENdy STATE PARK, VI-23-1973 / F.G. ANdREws, A.R. HARdy, M.S. WAsbAuER COLLECTORS (3Ƌ, 8♀ CSCA); MERCED CO., LOS BANOS VALLEY, 800', III-26-1 987, F. ANDREwS, A. HARDY & T. EICHLIN, IN STREAM (3Ƌ, 3♀ CSCA); MONTEREY CO., SANTA LUCIA, ARROYO SECO CAMP., V-2-64 / R.R. PINGER, JR. ACC. NO. 240 (1♀ CSCA); MONTEREY CO., STONE CANYON, CALIF., JAN. 1 1937, MENKE & STANGE / JON L. HERRING COLLECTION 1985 (1Ƌ USNM); NAPA CO., SAMUEL SPR., V-5-1955 / H.R. MOFFITT COLLECTOR (1♀ TAMU); NAPA CO., SAMUEL SPR., V- 18-1955 / R.W. BUSHING COLL. / C.J. DRAKE ACCESSION (1Ƌ USNM); SAN BENITO CO., PINNACLES NAT. MONUMENT, CHALONE CR., 15 JUNE 2002, COLL: R. WISSEMAN (1Ƌ, 1♀ UMC); SAN BENITO CO., PINNACLES NAT. MONUMENT, SANDY CR., 14 JUNE 2002, COLL: R. WISSEMAN (1♀ UMC); SAN B[ERNAR]D[I]NO CO., JUNE 15 1957, DEEP CREEK PUBLIC CAMP., CAL., MENKE & STAGE / C.J. DRAKE ACCESSION (2Ƌ, 1♀ USNM); SAME BUT / Ambrysus californicus MONT. dET. A. MENKE (1Ƌ USNM); SAN LUIS OBISPO CO., CHOLAME CREEK 1 MI. S. CHOLAME / 1.V.1963 H.B. LEECH & P.H. ARNAud, JR. cOLLEcTORs / Ambrysus californicus bohartorum USINGER DET. J.T. POLHEMUS 1968 (1Ƌ USNM); SAN LUIS ObIspO CO., ChOLAmE CREEK 1 mI S. ChOLAmE / 1-V-1963 / H.B. LEEch, P.H. ARNAud, JR. COLLEcTORs / Ambrysus californicus bohartorum DET. J.T. POLHEMUS (1♀ USNM); SAN LUIS OBISPO CO., CHOLAME CREEK 1 MI. S. CHOLAME / 1.V.1963 H.B. LEECH & P.H. ARNAUD, JR. COLLECTORS / C.J. DRAKE ACCESSION (1Ƌ, 1♀ USNM); SAME BUT / Ambrysus californicus californicus MONT. DET. J.T. POLHEMUS (1Ƌ USNM); S[ANTA] BARBARA CO., SANTA YNEZ RIVER, SAN LUCAS BGE. / 1-2-1939 CAL. / B.E. WhITE cOLLEcTOR / BuRdETTE E. WhITE COLLEcTION CALIf. AcAd. ScI. AccEssION 1967 / C.J. DRAKE ACCESSION (2♀ USNM); SAME BUT / Ambrysus californicus californicus MONT. dET. J.T. POLhEmus / C.J. DRAKE ACCESSION (1Ƌ USNM); [SANTA BARBARA CO.], BUELLTON CAL., 21 OCT. 1952, J.T. POLHEMUS / Ambrysus californicus MONTANdON 1897 (IRA LA RIvERs) / J. C. LuTz COLLEcTION 1961 / Ambrysus californicus USINGER (1Ƌ USNM); [SANTA BARBARA CO.], BUELLTON CAL., 21 OCT. 1952, J.T. POLHEMUS / C.J. DRAKE ACCESSION (2Ƌ, 3♀ USNM); SAME BUT / Ambrysus californicus MONTANDON DET. J.T. POLHEMUS / C.J. DRAKE ACCESSION (1♀ USNM); [SANTA BARBARA CO.], BuELLTON, CAL., 21 OcT. 1952, J.T. POLhEmus / Ambrysus californicus DET. J.T. POLHEMUS (1Ƌ, 1♀ USNM); SONOMA CO., GUALALA RIVER @ HAUPT CREEK, MAY 18, 2000, G.L. CHALLETT (1Ƌ TAMU); SONOMA CO., HAUPT CREEK AT GuALALA

RIVER, MAY 18, 2000, G.L. CHALLETT (1Ƌ, 1♀ TAMU); STANISLAUS CO., DEL PUERTO CAN., DEL PUERTO R. MP 16, 20 APRIL 2000, J.S. ASCHER COLR. (1Ƌ UMC); STANISLAUS CO., DEL PUERTO CAN., DEL PUERTO R., 21 APRIL 2000, J.S. ASCHER COLR. (1Ƌ, 5♀ UMC); STANISLAUS CO., DEL PUERTO CANYON, 3 MI. NW HIGHwAY 5, V-16-1987, F. ANDREwS & A. HARDY, UNDER LOG ALONG STREAM (1Ƌ CSCA); VENTURA CO., HOwARD CK. AT BEAVER CMPGRD., 24 NOV. 1990, COLL: J.A. BACK, AqUATIC VEGETATION (3Ƌ, 4♀ UMC); 6 MI. E. PT. REYES, CAL., IV-2-55 / S.W. HITCHCOCK COLLECTOR (2Ƌ, 1♀ USNM).