Nothogreniera occidentalis

A. Craig, Douglas, Currie, Douglas C. & Moulton, John K., 2018, Nothogreniera new genus, for two species of Australian " Paracnephia " (Diptera: Simuliidae), Zootaxa 4444 (4), pp. 351-380 : 368-378

Nothogreniera occidentalis ( Mackerras & Mackerras 1949, attrib. Tonnoir). New status and new combination

( FIgS. 53–83 )

' Simulium fergusoni occidentalis' Tonnoir. Unpublished manuscript name (nec Simulium occidentale Townsend 1891: 106 ; Smart 1945: 510). Nomen nudum.

Cnephia fergusoni (Tonnoir) var. Mackerras & Mackerras 1949: 386. (Type missing).

“ Cnephia ” ‘S. x. (west)’. Moulton 2000: 98. 2003: 43. Molecular examination.

Nothogreniera occidentalis (Mackerras & Mackerras) . New status and combination. The authors are following the precedent set by Crosskey & Howard (1997), who recognized Mackerras & Mackerras, 1949 as the authors of occidentalis . Curiously, Crosskey & Howard (loc cit.) did not comment on this muddy nomenclature.

Description. Adult female: (BASED ON FOUR PINNED PARATyPES, ONE SlIDE MOUNT ( FIg. 54 ) AND A PHARATE SPECIMEN). Body ( FIg. 53 ): TOTAl lENgTH 2.1–2.8 MM; MARKEDly BlACKISH gREy WITH POllINOSITy ON THORAX, lEgS BROWN. Head ( FIg. 55 ): OVERAll BlACK, POllINOSE IN SOME lIgHTINg, FRONS TAPERED ONly SlIgHTly TOWARDS ANTENNAE, NARROWEST JUST ABOVE ANTENNAE; WIDTH 0.6–1.1 MM; DEPTH 0.50–0.86 MM; POSTOCCIPUT VESTITURE OF SPARSE, SHORT BlACK HAIRS; FRONS BlACKISH BROWN; FRONS/HEAD WIDTH RATIO 1.0:3.8–4.6. Eyes: INTEROCUlAR DISTANCE 0.09–0.14 MM; OMMATIDIA DIAMETER 0.017 MM; ca. 27 ROWS ACROSS AND 35 DOWN AT MID-EyE. Clypeus: WIDTH 0.25–0.27 MM; DARK BROWN, POllINOSE; VESTITURE OF SPARSE PAlE HAIRS. Antenna ( FIg. 56 ): TOTAl lENgTH 0.6–0.9 MM; NINE ANTENNOMERES; TAPERED; SCAPE SMAll, PEDICEl MARgINAlly BROADER THAN ANTENNOMERE I, THAT AS BROAD AS lONg, REMAINDER gRADUAlly INCREASINg IN lENgTH, OCCASIONAl ANTENNOMERE MAlFORMED AND FUSED TO ANOTHER, DISTAl ANTENNOMERE SHORT, BUT lONgER THAN BROAD. Mouthparts: SUBSTANTIAl, ca. 0.6X lENgTH OF HEAD DEPTH; CIBARIUM MEDIAN DEPRESSION SHAllOW, SMOOTH WITH IRREgUlAR PIgMENTATION, CORNUAE SHORT, SUBSTANTIAlly SClEROTISED ( FIg. 57 ); MANDIBlE MARKEDly EXPANDED DISTAlly, ca. 16 OUTER AND 40 INNER TEETH, SMAll, DECREASED IN SIZE PROXIMAlly; lACINIA WITH 16–18 AND 17–19 TEETH ON INNER AND OUTER SIDES RESPECTIVEly ( FIg. 59 ); MAXIllARy PAlP ( FIg. 58 ), TOTAl lENgTH 0.66–0.69 MM, PAlPOMERES I AND II SMAll, III DARKER BROWN THAN REMAINDER, SlIgHTly EXTENDED BEyOND ARTICUlATION WITH IV, WHICH IS lIgHT BROWN AND SHORT, V (DISTAl) PAlE AND MODERATEly lONg; PROPORTIONAl lENgTHS III—V PAlPOMERES 1.0:0.5:0.9; SENSORy ORgAN NOT MARKEDly ElONgATED, IRREgUlAR OR OVAl (VARIABlE), 0.3–0.4X lENgTH OF PAlPOMERE III, OPENINg 0.4X VESIClE WIDTH. Thorax: lENgTH 1.1–1.4 MM; WIDTH 0.9 MM; POSTPRONOTAl lOBE CONCOlOUROUS WITH SCUTUM, MARKEDly HIRSUTE WITH FINE HAIR lONgER THAN THAT ON SCUTUM, SCUTUM OVERAll BlACK WITH EVEN SPARSE FINE gOlDEN HAIRS; SCUTEllAR DEPRESSION CONCOlOUROUS, BUT POllINOSE WITH lONgER SPARSE gOlDEN HAIRS; SCUTEllUM CONCOlOUROUS WITH SCUTUM, VESTITURE OF SPARSE MARKEDly lONg FINE gOlDEN HAIRS; ANTEPRONOTAl lOBE WITH DENSE lONg gREyISH HAIRS; PROEPISTERNUM WITH DENSE lONg PAlE HAIRS; FORE COXA HAIRED; PlEURON AND ANEPISTERNAl MEMBRANE DARK BROWN, lACKINg HAIRS; KATEPISTERNAl SUlCUS WEll EXPRESSED; METATHORACIC FURCASTERNUM lATERAl FlANgES NOT MARKEDly DEVElOPED ( AS FOR N. fergusoni , FIg. 8 ). Wing: ( FIg. 63 ) lENgTH 3.1–3.5 MM; WIDTH 1.3–1.5 MM; COSTA lACKINg SPINIFORM SETAE, HAIRlIKE SETAE ONly, R s WITH HAIRlIKE SETAE ONly; BASAl RADIAl CEll WEll DEVElOPED; A/B RATIO 1.0:2.7; M1 VEIN DOUBlE. Haltere: DARK yEllOWISH TO BlACKISH BROWN. Legs ( FIg. 60 ): EVENly DARK BROWN; HIND BASITARSUS WITH ROW OF WIDEly SPACED STOUT SPINES; CAlCIPAlA MARKEDly POORly DEVElOPED, WITH A FEW STOUT SPINES CONCENTRATED IN THAT POSITION, DISTINCT INTERSEgMENTAl PlATE VENTRAlly BETWEEN BASITARSUS AND TARSOMERE II, HIRSUTE, AlSO BETWEEN TARSOMERES II & III; PEDISUlCUS ABSENT; ClAWS SMAll, THUMBlIKE lOBE MARKEDly SMAll TO ESSENTIAlly ABSENT—VARIABlE, SMAll ROUNDED HEEl ( FIg. 61 ). Abdomen ( FIg. 62 ): ABDOMINAl SCAlE DARK BROWN, DUll yEllOWISH MEDIAlly WITH lONg CREAMy-yEllOW HAIRS EXTENDED TO ABDOMINAl SEgMENT III; REMAININg SEgMENTS EVENly DARK BROWN, WITH MOTTlED yEllOWISH AREAS DORSAlly; TERgITE II BROAD, 3.3X BROAD AS lONg, TERgITE III QUADRATIC WITH ROUNDED CORNERS, TERgITE IV SMAllER AND QUADRATIC, TERgITE V THE SMAllEST, VI lARgER AND TERgITE VII BROAD, VESTITURE ESSENTIAlly ABSENT ON ANTERIOR TERgITES, INCREASED POSTERIORly; PlEURITES POORly DEVElOPED; STERNITES ABSENT. Genitalia: STERNITE VIII EVENly PIgMENTED VESTITURE OF MICROTRICHIA AND SPARSE SUBSTANTIAl SETAE; HyPOgyNIAl VAlVES WITH MEDIAl EDgES STRENgTHENED AND STRAIgHT, gAP SlIgHTly TAPERED ANTERIORly, VESTITURE OF MICROTRICHIA AND SPARSE SETAE ( FIg. 64 ); gENITAl FORK WITH ANTERIOR ARM NARROWED AND CURVED DORSAlly (DISTORTED IN IMAgE), ANTERIOR APEX BAREly EXPANDED, lATERAl ARMS POORly SClEROTIZED, APODEME APPARENT, BUT WEll lATERAl, POSTEROlATERAl EXPANSIONS ROUNDED ( FIg. 65 ); CERCI BlUNTly ROUNDED, STRAIgHT DORSAl MARgIN, ANAl lOBE POORly DEVElOPED ( FIg. 67 ); SPERMATHECA OVOID, DARK BROWN, EXTERNAlly SMOOTH, INTERNAl FINE SPINES (AKA ACANTHAE) SPARSE AND NOT EASIly APPARENT EVEN AT HIgH MAgNIFICATION, ClEAR REgION SURROUNDINg JUNCTION WITH SPERMATHECAl DUCT BAREly EVIDENT, POORly DEVElOPED SCUlPTINg ( FIg. 66 ).

Adult male ( REARED SPECIMEN IN POOR CONDITION AND TWO PHARATE SPECIMENS). Body: IN ETHANOl, OVERAll COlOUR BlACK; TOTAl lENgTH 3.7–3.9 MM. Head : WIDTH 0.97 MM; DEPTH 0.92 MM. Eyes : UPPER OMMATIDIA, DIAMETER 0.030 MM, ca. 19 ACROSS AND 22 DOWN ; lOWER OMMATIDIA SMAll, DIAMETER 0.015 MM. Antenna: MARKEDly NARROWED, TOTAl lENgTH 0.76 MM; OVERAll BlACK, PEDICEl SMAll, SCAPE TWICE AS lONg AND ROUNDED, ANTENNOMERE I SlIgHTly NARROWER THAN SCAPE, REMAINDER TAPERED SMOOTHly TO SMAll APICAl ANTENNOMERE. Mouthparts: MARKEDly INSUBSTANTIAl; lENgTH 0.26X HEAD DEPTH ; MAXIllARy PAlP 0.74 MM lONg, PAlPOMERES I AND II SMAll, PAlPOMERE III EXTENDED SlIgHTly PAST JUNCTION WITH PAlPOMERE IV; PROPORTIONAl lENgTHS OF III–V PAlPOMERES 1.0:0.6:0.7, SENSORy VESIClE IRREgUlAR SPHERICAl SHAPE, OCCUPyINg 0.3X WIDTH OF PAlPOMERE III. Thorax: ONly MODERATEly DOMED, lENgTH 1.4 MM; WIDTH 1.0 MM; PRONOTAl lOBE, ANTEPRONOTAl lOBE AND PROEPISTERNUM HAIRED; SCUTUM EVENly VElVETy BlACK, VESTITURE OF SPARSE SHORT FINE PAlE HAIRS, DENSE AND lONg IN SCUTEllAR DEPRESSION; DENSE AND lONgER ON POSTPRONOTAl lOBE; SCUTEllUM CONCOlOUROUS WITH SCUTUM, BARE MEDIAlly, WITH SPARSE lONg PAlE HAIRS lATERAlly; POSTNOTUM CONCOlOUROUS WITH SCUTUM; PlEURAE DARK BROWN, MEMBRANE lACKINg HAIRS. Wing: DETAIlS NOT AVAIlABlE. Haltere : BASE OF STEM BlACK, KNOB DARK TAN. Legs: DETAIlS NOT AVAIlABlE. Abdomen: DETAIlS NOT AVAIlABlE. Genitalia: HEAVIly PIgMENTED; gONOCOXA WIDER THAN lONg, VESTITURE OF SPARSE HAIRS; gONOSTylUS BROAD, CURVED ON OUTER EDgE, TWO SUBSTANTIAl SHORT, BlUNT, APICAl SPINES (FIg. 68); VENTRAl PlATE COMPlEX (FIg. 69), 1.3X WIDER THAN lENgTH, BROADly ROUNDED POSTERIORly AND EXPANDED, NARROWED MEDIAlly, VESTITURE OF DISTINCT HAIRS; SMAll CENTRAl CARINA; BASAl ARMS BROAD, PARAMERE CONNECTOR WEll DEVElOPED; MEDIAN SClERITE APPARENTly NOT MARKEDly DEVElOPED; PARAMERE BlADE-lIKE, lIgHTly STRENgTHENED AlONg MEDIAl EDgE, ARTICUlATION WITH THE VENTRAl PlATE BASAl ARM EXTENSION NOT DISTINCT, BUT IS SO WITH THE gONOCOXAl APODEME, NO APPARENT TERMINAl SPINES; AEDEAgAl MEMBRANE WITH SMAll SPINES.

Pupa (BASED ON THREE IMMATURE SPECIMENS AND ONE EXUVIAE). BODy lENgTH ca. 3.0 MM ( FIg. 70 ). CUTIClE COlOURlESS AND THIN. FEMAlE HEAD CUTIClE BROAD BASAlly, lENgTH 1.5X BASAl WIDTH. SETAE NOT APPARENT ( FIg. 72 ). MAlE HEAD CUTIClE, UNKNOWN. THORACIC CUTIClE SMOOTH TO SlIgHTly RUgOSE ( FIg. 71 ), THORACIC DORSOCENTRAl SETAE ElONgATE, WITH SlIgHTly CURlED TIPS. GIll: WITH SHORT BASE, BRANCHINg INTO THREE; DORSAl AND MEDIAN BRANCHES SUBSTANTIAl WITH MUlTIPlE BRANCHINg POINTS, 15 lONg, lIgHT BROWN, MARKEDly FINE FIlAMENTS, lENgTH ca. 3.8 MM, SUBEQUAl TO PUPAl lENgTH. SURFACE STRUCTURE FINEly PSEUDOANNUlATED. ABDOMINAl ARMATURE AND PlEURITES AS FOR N. fergusoni ( SEE FIg. 36 ) ; ARMATURE EVEN lESS WEll DEVElOPED. Cocoon. APPARENTly JUST A FEW THREADS OF SIlK OVER THE POSTERIOR OF THE PUPAl ABDOMEN.

Larva (BASED ON FIVE lAST INSTAR lARVAE). Body (FIg. 73): lENgTH 5.8–6.7 MM. Head ( FIg. 74 ): lENgTH 0.8 MM, WIDTH 0.6 MM, COlOUR lIgHT yEllOW AND BROWN, VARIABlE, SOMETIMES COlOURlESS; HEAD SPOTS POSITIVE. EARlIER INSTAR HEAD MEDIUM BROWN. ANTEROMEDIAN AND POSTEROMEDIAl HEAD SPOTS POSITIVE AND DISTINCT, lATERAl SPOTS lESS SO; ECDySIAl lINES SINUOUS; CERVICAl SClERITES ATTACHED lATERAlly TO POSTOCCIPUT. Antenna ( FIg. 75 ): EXTENDED BEyOND FAN STEM, 0.6 MM lONg, DISTAl ANTENNOMERE ElONgATED AND SUBEQUAl TO MEDIAN PlUS BASAl ANTENNOMERE, PROPORTIONS 1.0:0.7:1.6. DISTAl ANTENNOMERE WITH MARKEDly FINE SClEROTIZED BANDS ( FIg. 75 ; INSERT). Labral fan: STEMS COlOURlESS, FANS MARKEDly DEVElOPED, ca. 70 FINE RAyS, 0.75–0.90 MM lONg, MID WIDTH ca. 0.006 MM, lONg MICROTRICHIA SUBEQUAl TO RAy WIDTH, INTERSPERSED WITH 4 OR 5 SMAllER. Maxilla ( FIg. 76 ): NOT HEAVIly PIgMENTED. Mandible ( FIg. 77 ): ElEgANTly CURVED AND ElONgATE, NOT MARKEDly PIgMENTED, ANTERIOR BRUSH WEll EXPRESSED, APICAl TOOTH WEll EXPRESSED AND PROTRUDED, PREAPICAl TOOTH HAlF SIZE OF APICAl TOOTH, SPINOUS TEETH MARKEDly FINE AND ElONgATED, SERRATIONS AND SENSIllUM DISTINCT AND WEll EXPRESSED, BlADE REgION FlAT ( FIg. 78 ). Postgenal cleft ( FIg. 79 ): POORly EXPRESSED, SHAllOW V-SHAPE WITH IRREgUlAR ANTERIOR APEX, DISTINCTly AND BROADly PIgMENTED AlONg EDgE, POSTgENA ESSENTIAlly COlOURlESS, PROPORTIONS OF HyPOSTOMA TO gENAE AND ClEFT 1.0:1.8:0.4. Hypostoma : TEETH MARKEDly PIgMENTED, IN THREE UNITS, TEETH 0, 1, 2 MARKEDly VARIABlE, TOOTH 0 EITHER NOT PROTRUDED OR DIRECTED DORSAlly (APPEARS ROUNDED AND FORESHORTENED IN VENTRAl VIEW, e.g. FIg. 79 ) WITH TEETH 1 AND 2 SMAll, SUBEQUAl IN SIZE, OR ( FIg. 80 ), TOOTH 0 DISTINCT WITH TOOTH 1 lATERAl ON BASE, TOOTH 3 SMAll ON BASE OF ENlARgED AND PROTRUDINg TOOTH 4, TEETH 5 & 6 lATERAlly ON BASE OF TOOTH 4, VARIABlE AND OCCASIONAlly POORly EXPRESSED, TOOTH 7 SMAll, TOOTH 8 EITHER NOT EXPRESSED, OR POORly SO ON ONE SIDE ONly; lATERAl SERRATIONS DOWN CONVEX SlOPED SIDES OF HyPOSTOMA SMAll, BUT DISTINCT. ONly ONE SUBSTANTIAl HyPOSTOMAl SETA WITH TWO POORly EXPRESSED OTHER SETAE MORE PROXIMAlly, VARIABlE. Anterior proleg ( FIg. 81 ): HOOKS MARKEDly FEW, lATERAl SClERITE POORly SClEROTIZED AND D-SHAPED. Thorax ( FIg. 82 ): PHARATE PUPAl gIll HISTOBlAST WITH FIlAMENT BASES CURVED VENTRAlly, TURNINg SMOOTHly POSTERIORly AND THEN DORSAlly AlMOST TO DORSAl ORIgIN OF gIll, THEN BENDINg RAPIDly VENTRAlly TO RECURVED DORSAlly AgAIN. Abdomen: MOTTlED gREyISH yEllOW, lIgHTER POSTERIORly; SOMETIMES WITH DORSAl SPOT PAIRS DORSAlly ON SEgMENTS V–VII, EXPANDED SlIgHTly TO SEgMENTS V–VII THEN DECREASED SMOOTHly TO POSTERIOR PROlEg. Ventral tubercles: ABSENT OR POORly EXPRESSED. Rectal papillae: THREE SIMPlE lOBES. Anal sclerite ( FIg. 83 ): WITH DORSAl AND VENTRAl ARMS SUBEQUAl IN lENgTH, MEDIAN REgION NOT MARKEDly DEVElOPED. Posterior circlet: 50–54 ROWS OF HOOKS, 10 OR 11 HOOKS PER ROW (TOTAl ca. 550).

Types. THE TAXONOMIC HISTORy OF N. occidentalis IS SOMEWHAT MURKy. ACCORDINg TO MACKERRAS & MACKERRAS (1949: 386), TONNOIR INTENDED TO DESCRIBE THIS ENTITy AS " Simulium fergusoni occidentalis ", EVEN DESIgNATINg A HOlOTyPE THAT SUPPOSEDly WAS DEPOSITED IN THE SCHOOl OF PUBlIC HEAlTH AND TROPICAl MEDICINE, SyDNEy . TONNOIR DIED IN 1940 ( MIllER 1940: 37) AND THIS DESCRIPTION WAS NEVER FORMAlly PUBlISHED. THE PUTATIVE ‘HOlOTyPE’ IS MISSINg AND CONSIDERABlE ATTEMPTS By DAC TO lOCATE IT HAVE PROVEN UNSUCCESSFUl. HOWEVER, ‘PARATyPES’, APPARENTly lABElED AS SUCH By IAN MACKERRAS, ARE NOW HOUSED IN ANIC, CANBERRA.

Nothogreniera occidentalis POSSESSED SUFFICIENT MORPHOlOgICAl DIVERgENCE FROM N. fergusoni TO BE CONSIDERED A VAlID SUBSPECIES By TONNOIR—WE CONCUR, AlTHOUgH AT FUll SPECIES lEVEl.

Holotype. WHEREABOUTS CURRENTly UNKNOWN, POSSIBly lOST; HOWEVER, IT MAy EVENTUAlly BE DISCOVERED IN AN AUSTRAlIAN COllECTION. ACCORDINgly, AT THIS TIME, WE REFRAIN FROM DESIgNATINg A NEOTyPE FROM ONE OF THE AVAIlABlE PARATyPES.

Paratypes. PINNED FEMAlES. LABEl DATA:- [ONE HIND lEg MOUNTED BETWEEN COVER SlIPS BITS] [BRIDgETOWN, W. A./ 29 AUg. 1926./ E. W. FERgUSON] [PARATYPE. (PRINTED)/ SIMUlIUM / FERgUSONI/ OCCIDENTAlIS/ TONN (BlUE, HANDWRITTEN)] [ CNEPHIA / FERgUSONI VAR./ DET. MACKERRAS] [ANIC DATABASE NO./29 029386] [AUST. NAT./ INS. COLL. (gREEN)].

[BRIDgETOWN W.A./ 29 AUg. 1926 / E. W. FERgUSON] [PARATYPE/ SIMUlIUM / FERgUSONI/ OCCIDENTAlIS/ TONN] [ANIC DATABASE NO./ 29 029385] [AUST. NAT./ INS. COll.], PlUS NO./ 29 0 29387 AND NO. 29 029382—NOW MOUNTED ON SlIDE.

A PINNED SPECIMEN (NO. K67560 ), INDICATED AS A PARATyPE, IS HOUSED IN THE AUSTRAlIAN NATIONAl MUSEUM, SyDNEy. COllECTED By E. W. FERgUSON. DATA gIVEN IS, IN PART— "PARATyPE: SIMUlIUM FERgUSONI OCCIDENTAlIS TONNOIR, MSS NAME: SIMUlIIDAE : DIPTERA : INSECTA: …….”. THE DATE OF COllECTION WAS 31-VIII-1926, TAMMIN, WA (S31.6400° E117.4800°). IT IS NOT ClEAR IF THIS SPECIMEN IS STRICTly A PARATyPE. IT IS lISTED UNDER " Paracnephia fergusoni " IN THE “ATlAS OF LIVINg AUSTRAlIA” <HTTPS://BIOCACHE.AlA.ORg.AU/OCCURRENCES/ SEARCH?Q=lSID:URN:lSID:BIODIVERSITy.ORg.AU:AFD.TAXON:ED160F9B-A369-4D87-9EDD-F5B1B298C009#TAB_RECORDSVIEW>. SIMIlARly, A SPECIMEN ‘K179806’, 30-VIII-1926, NARROgIN, WA (S32.9300° E117.1700°).

Additional material. PINNED FEMAlES:- [3M. S. MT. COOKE/ WA./ 30.VIII.56 / O/L/ MCINTOSH] [ CNEPHIA / FERgUSONI/ VAR. (TONN.)/ I. M. MACKERRAS DET.] [AUST. NAT./ INS. COll.] AND [TAMMIN/ 31 AUg ‘26/ NICHOlSON] [WESTERN/ AUSTRAlIA] [ CNEPHIA / FERgUSONI/ VAR./ DET MACKERRAS] [AUST. NAT./ INS. COll.]. SlIDE MATERIAl:- FEMAlE [ANIC DATABASE NO/ 29 029359] [ CNEPHIA / FERgUSONI VAR./ {F}/ BRIDgETOWN W.A./ 29/8/26. A.J.N]; ON BACK OF SlIDE [AUST. NAT./ INS. COll.] ( FIg. 54 ); FEMAlE [ Nothogreniera / occidentalis ] [WESTERN AUSTRAlIA/ DAlE RIVER TRIBUTARy/ S32° 21’ 30”/ E 116° 27’ 30”/ ElEV. 300M. 7-IX-1996 / COll. JK MOUlTON] [ANIC DATABASE NO. 29 026653]. LAST INSTAR lARVA, AS ABOVE [ANIC DATABASE NO. 29 026652]. All STAgES (UASM# 353138–353145). AlCOHOl MATERIAl:- TWO lARVAE AND A PUPA. LABEl DATA:- [ Nothogreniera / occidentalis ] [AUSTRALIA, WA/ LAST STREAM ON N. METRO RD./ BEFORE X-INg WITH MCCAllUM/ RD. TRIB. OF DAlE RIVER/ S32° 21’ 30”/ E 116° 27’ 30”/ 7 SEPTEMBER 1996 / COll. J. K. MOUlTON] [ANIC DATABASE NO. 29 026651]. LARVAE, PUPAE, REARED MAlE:- [DAlE R. TRIB (11-5)/ 11-VII-81 JP (AKA JANE PRINCE)] (UASM# 353367) AND [AUSTRALIA, WA/ LAST STREAM ON N. METRO RD./ BEFORE X-INg WITH MCCAllUM/ RD. TRIB. OF DAlE RIVER/ S32° 21’ 30”/ E 116° 27’ 30”/ 7 SEPTEMBER 1996 / COll. J. K. MOUlTON], (UASM# 353365, 353366). THE PUPAE ARE OF VERy EARly STAgE AND SUBOPTIMAl FOR DESCRIPTION.

Etymology. ORIgINAlly 'OCCIDENTAlIS' WAS AN UNPUBlISHED SUBSPECIES MANUSCRIPT NAME By TONNOIR ( MACKERRAS & MACKERRAS 1949: 386), AlBEIT WRITTEN ON SPECIMEN lABElS. WHIlE NOT USED By MACKERRAS & MACKERRAS (loc. cit.), WE DO SO HERE FOR HISTORICAl VAlUE.

Distribution ( FIg. 84 ). Southern Western Australia: MT. COOKE, S32.3500° E116.3300°, ElEV. 300M., 30-VIII- 1956 (ANIC); BRIDgETOWN, S33.9700° E116.1400°, ElEV. 200M., AUgUST ( MACKERRAS & MACKERRAS 1949); NARROgIN, S32.9300 E117.1800, ElEV. 360M., AUgUST ( MACKERRAS & MACKERRAS 1949); TAMMIN, S31.6400° E117.4800°, ElEV. 240M., 31-VIII-1926 ( MACKERRAS & MACKERRAS 1949); DAlE RIVER TRIBUTARy, S32.3586° E116.4583°, ElEV. 300M., 7- IX-1996 (JKM); DAlE RIVER TRIBUTARy, S32.3500° E116.4500°, ElEV. 300M., 8-IX-1996 (JKM); DRyANDRA, S32.7200° E116.9300°, ElEV. 300M., 31-VIII-1956 (ANIC); DONNyBROOK, S33.5900° E115.8200°, ElEV. 80M., 1-IX-1952 (ANIC).

Bionomics. All ORIgINAl FEMAlE ADUlTS WERE COllECTED IN AUgUST. IT CAN BE ASSUMED THAT THEy WERE NETTED WHIlE ATTEMPTINg TO BITE THE COllECTOR. NOTHINg ElSE IS KNOWN. THE IMMATURE STAgES COllECTED By JKM WERE TAKEN FROM A SMAll TRIBUTARy OF THE DAlE RIVER ON THE DARlINg ESCARPMENT, WESTERN AUSTRAlIA IN SEPTEMBER. THE lOCAlITy WAS JUST A SANDy DITCH WITH lOW FlOW AND A FEW SUBSTRATES OF STICKS, lEAF PACKS, AND EVEN RARER TRAIlINg gRASS—IN FUll AgREEMENT WITH THE PHySIOgNOMy OF THE lARVAE. LARgE POROUS lABRAl FANS HAVE BEEN SUggESTED ( PAlMER & CRAIg 2000; AND OTHERS) AS AN ADAPTATION TO SlOW FlOW AND HIgH SESTON lEVElS, AND THERE IS THE WEll KNOWN CORRElATION (e.g. PAlMER & CRAIg loc. cit.; KONURBAyEV 1973; AND AgAIN, OTHERS) BETWEEN lOW VElOCITy AND SMAll NUMBERS OF HOOKS IN THE POSTERIOR CIRClET—FOR N. occidentalis SOME 550. WHIlE AMONgST THE lOWEST NUMBERS RECORDED By CROSSKEy (1990: 163) AND PAlMER & CRAIg (loc. cit.), " Paracnephia " umbratora lARVAE POSSESS A MERE 390 HOOKS. SIMIlAR lOW NUMBERS ARE KNOWN FOR Bunyipellum gladiator AND EVEN lOWER FOR UNDESCRIBED WESTERN AUSTRAlIA SPECIES RElATED TO " P. " umbratora .

THE lONg ANTENNAE ON lARVAE OF N. occidentalis AND, lESS SO THOSE OF N. fergusoni , POINT TO ADAPTATION TO THE SlOW FlOW OF THEIR KNOWN HABITATS. BOTH CROSSKEy (1990: 108) AND ADlER et al. (2004: 53) COMMENT THAT lONgER ANTENNAE ARE COMMON FOR SIMUlIID lARVAE INHABITINg SlOW FlOWS. THIS AgREES WITH A STUDy By CRAIg (1990: 349) ON THE EPHEMEROPTERAN Cloeon dipterum (LINNAEUS), WHERE lONg ANTENNAE ARE USED TO PENETRATE THE BOUNDARy lAyER SURROUNDINg THE BODy WHEN THE lARVA IS MOVINg SlOWly.

DATES OF COllECTION INDICATE THAT N. occidentalis IS A UNIVOlTINE AUSTRAl SPRINg SPECIES, SIMIlAR TO N. fergusoni .

Remarks. AS NOTED ABOVE, THE N. occidentalis HAS A SOMEWHAT CHECKERED TAXONOMIC HISTORy. TONNOIR PROPOSED THE NAME IN A MANUSCRIPT, BUT IT WAS NEVER FORMAlly PUBlISHED. THENCE THE ‘TyPES’ HE DESIgNATED AND NAMES ON MATERIAl STIll IN EXISTENCE, HAD NO VAlIDITy. HOWEVER, MACKERRAS & MACKERRAS (1949: 386) HAD ACCESS TO TONNOIR’S ‘PARATyPES’ AND DESCRIBED THE SPECIES AS “ Cnephia fergusoni (TONNOIR) VAR.” (e.g. FIg. 54 ). PRIOR TO 1960, THE ICZN AllOWED SUCH AS VAlID SUBSPECIES.

HERE WE FORMAlly RAISE THAT ENTITy TO SPECIES STATUS AND ASSIgN IT TO Nothogreniera AS N. occidentalis (MAKERRAS & MACKERRAS). IT IS SIMIlAR IN MANy RESPECTS TO N. fergusoni , WITH POSSESSION OF A ClUMP OF STIFF SPINES ON THE INNER APICAl MARgIN OF THE HIND BASITARSUS (i.e., IN THE VICINITy OF THE CAlCIPAlA, WHEN PRESENT), MINUTE TOOTH ON THE FEMAlE ClAW, STRENgTHENED INNER MARgINS OF THE HyPOgyNIAl VAlVES, STRIATIONS ON THE DISTAl ANTENNOMERE OF THE lARVAl ANTENNA AND THE MARKEDly TRIlOBED APPEARANCE OF THE HyPOSTOMAl TEETH. ADUlT N. occidentalis ARE OVERAll DISTINCTly MORE BlACKISH IN COlOUR, WITH ASHy REFlECTIONS ON THORAX, WITH lEgS lIgHTER BROWN (cf. FIgS. 1, 2 , 53 ). THE lARVAE HAVE lARgE lABRAl FANS, lONgER ANTENNAE, A MORE ACUTEly POINTED POSTgENAl ClEFT AND MORE yEllOWISH BODy COlOURATION. COllECTIVEly, THESE DIFFERENCES ARE CONSIDERED SUFFICIENTly DISTINCT TO WARRANT RECOgNITION OF N. occidentalis AS A VAlID SPECIES. INDEED, MOUlTON (2003: 50), IN A MOlECUlAR STUDy, SHOWED N. fergusoni AND N. occidentalis (AS “C.” ‘S.X. (EAST)’ AND “C.” ‘S.X. (WEST)’ RESPECTIVEly) TO BE SISTER SPECIES WITH HIgH SUPPORT.

Discussion. WITH MORE STAgES NOW AVAIlABlE, NAMEly THE IMMATURES, IT IS ClEAR THE AUSTRAlIAN SO-CAllED " Paracnephia " SPECIES DO NOT BElONg IN THAT AFRICAN gENUS AS ASSIgNED By CROSSKEy & HOWARD (1997).

Nothogreniera , WHIlE SHARINg SOME CHARACTER STATES WITH AFRICAN Paracnephia (SEE DE MOOR 2017: 712), DOES NOT FIT EASIly INTO ANy OF THE SPECIES-gROUPS OF THAT gENUS. THE BEST FIT WOUlD BE IN THE muspratti -gROUP ( CROSSKEy 1969). EVEN THERE, HOWEVER, AT MINIMUM, THE TERMINAl SPINES ON THE PUPAl ABDOMEN OF Nothogreniera ARE SHORT AND HOOKS ARE ABSENT FROM STERNITE IV. NO MENTION IS MADE By CROSSKEy (loc. cit.) OF EXTENDED ANTENNAE OF lARVAE IN ANy DESCRIPTIONS OF Paracnephia SPECIES, OR FOR THAT MATTER OF ANy PATTERNINg ON THE DISTAl ANTENNOMERES. THAT lATTER CHARACTER STATE ( FIgS. 41 , 75 ) IS SEEN IN SEgREgATES OF Greniera (ADlER et al. 2004: 280) AND IS CERTAINly UNIQUE IN AUSTRAlIAN SIMUlIIDS. NEITHER IS THERE ANy MENTION FOR Paracnephia s.s. OF THE AggREgATION OF STIFF SPINES ON THE CAlCIPAlA THAT TONNOIR (1925: 221, HIS FIg. 2 G ), IllUSTRATED FOR N. fergusoni . THIS SPINE ClUMP, AT THE TIME UNIQUE IN AUSTRAlIAN SIMUlIIDS, IS NOW KNOWN FOR Bunyipellum gladiator AND AlSO FOR UNDESCRIBED SPECIES RElATED TO Paracnephia umbratora .

FURTHER, IN SOME SPECIES OF THE muspratti -gROUP OF Paracnephia THE ADUlT FEMAlES lACK TEETH ON THE MANDIBlE—REPlACED WITH HAIRS, WITH MAlES POSSESSINg AN ARRAy OF TERMINAl SPINES ON THE gONOSTylUS; CHARACTER STATES MARKEDly AT VARIANCE TO THOSE IN Nothogreniera .

Nothogreniera IS MORPHOlOgICAlly PERHAPS ClOSER TO SOUTH AMERICAN Araucnephia OR Araucnephiodes AND IS CERTAINly OF THAT gRADE OF DEVElOPMENT. NONETHElESS, Nothogreniera DIFFERS IN A NUMBER OF CHARACTER STATES FROM THOSE TWO gENERA, WITH N. fergusoni ADUlTS NOT KEyINg OUT EASIly IN THOSE By OF WygODZINSKy & COSCARóN (1973: 141). MAlE gENITAlIA MATCH MODERATEly WEll WITH THOSE OF Araucnephia , IN PARTICUlAR THE FORM THE VENTRAl PlATE AND PARAMERES (WygODZINSKy & COSCARóN loc. cit.; PESSOA et al. 2012). LARVAE HAVE A SIMIlAR ARRANgEMENT OF HyPOSTOMAl TEETH BUT DIFFER IN THAT Nothogreniera lACKS RECTAl SCAlES

WygODZINSKy & COSCARóN (1973) COMPARED Araucnephia TO AUSTRAlIAN “ Cnephia ”, NAMEly strenua (MACKERRAS & MACKERRAS) AND tonnoiri (DRUMMOND) AND CONClUDED THAT THEy WERE NOT CO-gENERIC—WE AgREE. GIl- AZEVEDO (2010) IN A PHylOgENETIC ANAlySIS USINg MORPHOlOgICAl CHARACTER STATES, SHOWED (HIS FIg. 1 , PAgE 265) N. fergusoni + N. occidentalis AS SISTER TAXA, AlBEIT UNRESOlVED TO OTHER SOUTHERN HEMISPHERE gENERA. MOUlTON (2003: 47) REFERRED TO THESE TWO Nothogreniera SPECIES AS " CNEPHIA " 'S. X (EAST)' AND " CNEPHIA " 'S. X. (WEST)' RESPECTIVEly. IN HIS STRICT CONSENSUS ANAlySIS FROM THE 28S gENE, THESE ARE SISTER SPECIES, UNRESOlVED TO OTHER SOUTHERN HEMISPHERE SIMUlIIDS, IN AgREEMENT WITH GIl-AZEVEDO ABOVE. HOWEVER, A NEIgHBOUR JOININg ANAlySIS (HIS FIg. 2B ) USINg THE EF-1 α NUClEOTIDES SHOWED THE TWO SPECIES AS SISTER TO Greniera denaria + Mayacnephia (NOW Tlalocomyia ). Greniera IS A NORTHERN HEMISPHERE gENUS, AS IS ONE SEgREgATE OF Tlalocomyia , ANOTHER IN CENTRAl AMERICA (ADlER & CROSSKEy 2017: 120). THE PUTATIVE ClOSE RElATIONSHIP BETWEEN Nothogreniera , Greniera AND Tlalocomyia IS IN AgREEMENT WITH THE EXPRESSION OF THE lARVAl HyPOSTOMAl TEETH AND DETAIlS OF THE MAlE gENITAlIA (e. g., ADlER et al. 2004).

THE SPERMATHECA IS AN UNDERUTIlIZED SOURCE OF CHARACTERS IN PHylOgENETIC STUDIES OF SIMUlIIDS. EVANS & ADlER (2000) SURVEyED VARIATION IN A NUMBER OF CHARACTER STATES THROUgHOUT THE FAMIly, INClUDINg (AMONg OTHER FEATURES) THE DEgREE OF SClEROTIZATION AT THE JUNCTION OF THE SPERMATHECAl DUCT AND THE MAIN BODy OF THE SPERMATHECA, AND THE PRESENCE OR ABSENCE OF INTERNAl SPICUlES (ACANTHAE). A SPERMATHECA WITH AN UNPIgMENTED RINg SURROUNDINg THE JUNCTION OF THE DUCT WAS HyPOTHESIZED TO BE PlESIOMORPHIC—THE CONDITION FOUND IN Nothogreniera . EVOlUTIONARy PATTERNS OF INTERNAl SPICUlES ARE SOMEWHAT MORE COMPlEX, WITH EVANS AND ADlER (loc. cit.) HyPOTHESIZINg THE INDEPENDENT ACQUISITION OR lOSS OF SPICUlES IN CERTAIN TAXA. NONETHElESS, THE PRESENCE OF INTERNAl SPICUlES IS ARgUABly IN THE gROUND PlAN OF THE SIMUlIINI BASED ON THE PRESENCE OF THAT STATE IN Nothogreniera AND SEVERAl OTHER PRIMITIVE-gRADE SIMUlIINE gENERA INClUDINg Araucnephia, Araucnephioides, Cnesia, Cnesiamima, Gigantodax, Lutzsimulium , AND Paraustrosimulium .

ON BAlANCE, THE COMBINATION OF CHARACTERS STATES INDICATES THAT Nothogreniera SHOUlD BE CONSIDERED A SEPARATE gENUS. THE DEgREE OF EXPRESSION OF NUMBERS OF SUCH STATES (e.g., WINg VENATION, CAlCIPAlA EXPRESSION, lARVAl HyPOSTOMA ) INDICATES A PlESIOMORPHIC gRADE OF DEVElOPMENT.

THE EAST/WEST DISTRIBUTION OF SISTER SPECIES IS A COMMON PATTERN IN AUSTRAlIAN BIOTA (e. g., THOMPSON & STIllWEll, 2010) AND WEll EXEMPlIFIED FOR SIMUlIIDS (e.g., MACKERRAS & MACKERRAS 1949: 380, CRAIg et al. 2017, CRAIg et al. 2018). SUCH POINTS TO A VICARIANT EVENT OF GONDWANAN AgE—lIKEly THE SOUTHERN EXTENSION OF THE INlAND EROMANgA SEA THAT BROKE THROUgH CENTRAl-SOUTH AUSTRAlIA IN THE EARly CRETACEOUS (ca. 145 MyA), WHEN AUSTRAlIA WAS SEPARATINg FROM ANTARCTICA, AND WHICH lED TO THE FORMATION OF THE NUllABOR PlAINS, A MAJOR ARID BARRIER FOR AUSTRAlIA BIOTA. CERTAINly, SIMUlIIDAE WERE IN AUSTRAlIA THEN; FOSSIl lARVAE OF ClEAR SIMUlIID gESTAlT ARE KNOWN FROM THE CRETACEOUS APTIAN STAgE (116 MyA) ( JEll & DUNCAN, 1986, JEll 2004).

Figures

FIGURE 84. Map of southern part of Australia, showing distribution of Nothogreniera.

FIGURES 53–58. Nothogreniera occidentalis female. (53) Habitus. (Dryandra). Scale bar =1.0 mm. (54) An original slide mount of N. occidentalis by Tonnoir. (55) Cleared head. (Dale River). Scale bar = 0.2 mm. (56) Antenna. (Dale River). Scale bar = 0.1 mm. (57) Cibarium (Dale River). Scale bar = 0.1 mm. (58) Maxillary palp & lacinia. (Dale River). Scale bar = 0.1 mm.

FIGURES 59–63. Nothogreniera occidentalis female. (59) Mandible & lacinia. Scale bar = 0.02 mm. (60) Hind basitarsus. (Dale River). Arrow indicates concentration of spines on minute calcipala. Scale bar = 0.05 mm. (61) Claw, paratype. Arrow indicates minute tooth. Scale bar = 0.02 mm. (62) Tergites. (Dale River). Scale bar = 0.5 mm. (63) Wing, paratype. Scale bar = 0.5 mm.

FIGURES 64–67. Nothogreniera occidentalis female. FIGURES 68, 69. Male. (64) Hypogynial valves. (Dale River). Scale bar = 0.1 mm. (65) Genital fork. (Dale River). Scale bar = 0.05 mm. (66) Spermatheca. (Bridgetown). Scale bar = 0.05 mm. (67) Anal lobe & cercus. (Bridgetown). Scale bar = 0.05 mm. (68) Gonocoxa & gonostylus. gc—gonocoxa; gs—gonostylus. Scale bar = 0.05 mm. (69) Genitalia, pharate specimen. Stained. a m—adeagal membrane; b a—basal arm; p—paramere; p c—paramere connector; v p—ventral plate. Scale bar = 0.05 mm.

FIGURES 70–72. Nothogreniera occidentalis pupa (Dale River). FIGURES 73. Larva. (70) Habitus of male pupa. Scale bar = 1.0 mm. (71) Dorsal view of thorax, showing dorsocentral setae. Scale bar = 0.5 mm. (72) Female cephalic capsule. Scale bar = 0.5 mm. (73) Habitus, last instar larva. Scale bar = 1.0 mm.

FIGURES 74–79. Nothogreniera occidentalis, last instar larva. (74) Head. Scale bar = 0.5 mm. (75) Antenna. Scale bar = 0.1 mm. Insert, DIC of distal article. Scale bar = 0.02 mm. (76) Maxilla. Scale bar = 0.1 mm. (77) Mandible. Scale bar = 0.1 mm (78) Mandible tip. Scale bar = 0.05 mm. (79) Postgenal cleft & hypostoma. Scale bar = 0.2 mm.

FIGURES 80–83. Nothogreniera occidentalis last instar larva. (80) Hypostoma. Scale bar = 0.05 mm. (81) Anterior proleg, penultimate larva. Scale bar = 0.1 mm. (82) Lateral view of thorax, showing gill histoblast. Scale bar = 0.5 mm (83) Anal sclerite & circlet of hooks. Scale bar = 0.2 mm

FIGURES 7–11. Nothogreniera fergusoni adult female. (7) Maxillary palpus, lacinia. Scale bar = 0.2 mm. Insert, lacinia tip. Scale bar = 0.05 mm. (8) Mandible tip. Scale bar = 0.05 mm. (9) Cibarium, paratype (ANIC No. 29 029396. Bumberry). Scale bar = 0.1 mm. (10) Anepisternal membrane, paratype. (As for Fig. 9). Scale bar = 0.1 mm. (11) Wing, paratype. (As for Fig. 9). Scale bar = 0.5 mm.

FIGURES 34–38. Nothogreniera fergusoni, pupae (exuviae). (34) Cephalic capsule, female. Scale bar = 0.2 mm. (35) Cephalic capsule, male. Scale bar = 0.2 mm. (36) Abdomen, slide mount. Scale bar = 1.0 mm. (37) Thoracic cuticle. Scale bar = 0.2 mm. (38) Segment IX, showing terminal spines. Arrow indicates grapnel hooks. Scale bar = 0.1 mm.

FIGURES 1–6. Nothogreniera fergusoni adult female. (1) Habitus of holotype (Bumberry, 1916). Scale bar = 1.0 mm. (Image courtesy of Russell Cox, Australian Museum, Sydney). (2) Holotype labels. (Image courtesy of Melissa Graf, Australian Museum, Sydney). (3) Habitus of female. In ethanol. (Grampians National Park. 2014). Scale bar = 1.0 mm. (4) Paratype labels. (Australian National Insect Collection, Canberra). (5) Frontal view of head. (Grampians National Park). Distal palpomere foreshortened. Scale bar = 0.2 mm. (6) Antenna, paratype (ANIC No. 29 029396. Bumberry). Scale bar = 0.2 mm.

FIGURES 39–43. Nothogreniera fergusoni, last instar larva. (39) Habitus. Scale bar = 1.0 mm. (40) Dorsal head. Scale bar = 0.5mm. (41) Antenna. Scale bar = 0.05 mm. Insert: high magnification DIC of apical antennal article. Scale bar = 0.02 mm. (42) Mandible of last instar. Scale bar = 0.1 mm (43) Mandible tip. Scale bar = 0.02 mm.