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How to distinguish Neocallichirus, Sergio, Podocallichirus and Grynaminna (Decapoda: Callianassidae: Callichirinae) from each other in the fossil record?
0DW~ã+\åQê1 and Hiroaki Karasawa2
1'HSDUWPHQWRI*HRORJ\DQG3DODHRQWRORJ\)DFXOW\RI1DWXUDO6FLHQFHV&RPHQLXV8QLYHUVLW\0O\QVNi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“1HRFDOOLFKLUXV”JUDQGLV.DUDVDZDDQG*RGD IURPWKH0LGGOH3OHLVWRFHQHRI-DSDQLVUHYLVHGDQGDVVLJQHGWRWKHJHQXV*U\QDPLQQD .H\ZRUGV 'HFDSRGD&DOOLDQDVVLGDH1HRFDOOLFKLUXV6HUJLR3RGRFDOOLFKLUXV*U\QDPLQQDIRVVLOUHFRUGV\VWHPDWLFV ,QWURGXFWLRQ Generic assignment of callianassid ghost shrimps One of the greatest challenges of modern decapod crustacean 7KHELRORJLFDOFODVVL¿FDWLRQRIWKH&DOOLDQDVVLGDH'DQDLVEDVHG palaeontology is the interpretation, both systematic and ecological, mainly on soft part morphology, which include the dorsal carapace of callianassid burrowing ghost shrimps (Decapoda: Axiidea: architecture, the nature of maxillipeds, form of the abdomen, pleopods, Callianassidae). Their fossil record is very robust and they are present uropods and telson; for discussion on this issue with respect to the fossil in most associations of Cenozoic decapod crustaceans described so far. record see e.g. Schweitzer and Feldmann (2002) and Schweitzer et al. However, the generic assignment of the callianassid remains is very D 7KHFODVVL¿FDWLRQLWVHOILVKRZHYHUQRWFRQVLVWHQWWKURXJKRXW GLI¿FXOWSDUWO\EHFDXVHWKHUHLVJUHDWLQFRQVLVWHQF\DOVRLQWKHELRORJLFDO the biological literature and is in a need of comprehensive revision. FODVVL¿FDWLRQDQGWD[RQRP\RIWKLVJURXS 0DMRUPRQRJUDSKVIRFXVHGRQWKLVJURXSUHFHQWO\SXEOLVKHGE\6DNDL This contribution aims to discuss the generic assignment of several PHWOLWWOHDJUHHPHQWIURPWKHVLGHRIRWKHUZRUNHUV callianassid genera of the subfamily Callichirinae Manning and Felder, (Tudge et al 3RRUH 'ZRUVFKDN )HOGHUDQG 1991 in the fossil record when dealing with chelipeds only. In biological Robles, 2009). In general, there are several different views on the literature there is often little attention paid to the nature of chelipeds evaluation of taxonomically important characters as exemplified by when defining genera (but see the discussion on differences between ZRUNVRI%LIIDU 0DQQLQJDQG)HOGHU 3RRUH eucalliacine genera in Ngoc-Ho, 2003), which are usually the only fossil DQG6DNDL remains of these animals. In this respect assigning fossil material to the 0DQ\H[WDQWFDOOLDQDVVLGJHQHUDKDYHEHHQHUHFWHGZLWKRXWWDNLQJ genera Neocallichirus6DNDL Sergio Manning and Lemaitre, 1994; detailed account on their chelipeds. No wonder, because chelipeds in Podocallichirus6DNDL DQGGrynaminna Poore, 2000 is a very general are very variable features. There are many species with proven complex topic, as no general agreement about their systematic status is SRO\PRUSKLVPLQWKHLUQDWXUH HJ6DNDL LQVHYHUDOJHQHUD currently at hand. chelipeds are sexually dimorphic which is often connected with the DOORPHWULFJURZWK HJ)HOGHUDQG/RYHWW 'ZRUVFKDN On the other hand there are characters present on chelipeds which can EHFRQVLGHUHGRIWD[RQRPLFLPSRUWDQFH7KHZRUNRI%LIIDU FDQ be considered as first such step towards the systematic evaluation of 6RPHWLPHVWKHPDWHULDOPD\EHORQJWRDFORVHO\UHODWHGIDPLO\ VXFKFKDUDFWHUV+HSURYLGHGWZRVHSDUDWHNH\VWRWKH:HVWHUQ$WODQWLF Ctenochelidae sensu0DQQLQJDQG)HOGHU QRW6DNDL Callianassa species; one of them evaluates major chelipeds only. Later 2011). Manning and Felder (1991) emphasized the importance of some hard- $QDSSURDFKWU\LQJWR¿QGXQLI\LQJFKDUDFWHUVSUHVHQWRQFKHOLSHGV part morphology characters for generic assignment, some of them of taxa classified within a certain genus concept based on soft part SUHVHQWRQPHUXVRIPDMRUFKHOLSHG)ROORZLQJWKHLUZRUNELRORJLVWV morphology. In this approach extant congeneric taxa are studied in detail XVXDOO\DGGDSUHVHQFHRUDEVHQFHRIDPHUDOKRRNLQWKHGLDJQRVHVRI to identify these common characters on chelipeds. It is needed to test new genera or emendations of older ones. Unfortunately, it is of little different generic concepts to see whether presence of chosen characters help to palaeontologists, because there are many genera both with and is consistent throughout the supposedly congeneric taxa or not. This ZLWKRXWPHUDOKRRN%HFDXVHFKHOLSHGVDUHPRVWGXUDEOHSDUWVRIJKRVW DSSURDFKLVEDVLFDOO\DERXWORRNLQJIRUDJHQHULFGLDJQRVLVEDVHGRQ VKULPSVDQGDUHYHU\IUHTXHQWLQWKHIRVVLOUHFRUGSDODHRQWRORJLVWVFDQ chelipeds only which would not contradict the diagnosis based on soft often rely solely on the characters present on them. Therefore an affort part morphology. to identify diagnostic characters present not only on merus but also on other cheliped elements is needed. Neocallichirus in the fossil record ,QWKHZRUOGRISDODHRQWRORJ\WKHZRUNRI0DQQLQJDQG)HOGHU (1991) triggered several reassignments of fossil taxa previously referred The genus Neocallichirus is relatively widely defined and after to “Callianassa sensu lato”. Since then there is always an attempt to Callianassa it is the most speciose extant callianassid genus (De FODVVLI\IRVVLOFDOOLDQDVVLGVZLWKLQELRORJLFDOO\GH¿QHGJHQHUD Grave et al., 2009). For generic assignment always a combination of We discuss here basically two approaches of assignment of the fossil characters has to be used, as also such important taxonomic character material to respective callianassid genera: DVUGPD[LOOLSHGLVQRWVSHFL¿FVROHO\IRUNeocallichirus 6DNDL 1) An approach refusing to identify the genus on the hard part Manning and Felder (1991) reconsidered the diagnosis of Neocallichirus PRUSKRORJ\ FKHOLSHGVRQO\ VLPSO\EHFDXVHRIODFNRIVRIWSDUW and added also the characters on the chelipeds, notably on the merus. morphology characters on which the diagnoses of genera are usually The merus of Neocallichirus is variable in shape but is always serrated based. This approach may use approximate generic assignment as along the lower margin. According to Manning and Felder (1991) it “Callianassa” or Callianassa sensu latoDVH[HPSOLILHGE\ZRUNV ODFNVDPHUDOVSLQHRUKRRN DFFRUGLQJWR6DNDL LWLVZLWK of Müller (1984) and Karasawa (1998, 2000), respectively. If the RUZLWKRXWPHUDOKRRN PDLQO\GXHWRWKHV\QRQ\PLVDWLRQZLWKWKH fossil material fails to be assigned more closely, it is recommended genus Sergio ZKHUHDV6DNDL VWDWHGWKDWPHUXVLVZLWKRUZLWKRXW to classify fossil specimens within Callianassidae (or Callianassa) ventral convexity. Schweitzer and Feldmann (2002) and Schweitzer et sensu lato without reference to a subfamily or genus (Collins et al., al D GLVFXVVHGXVDJHRIWKHFKDUDFWHUVRQPDMRUFKHOLSHG QRW 7DEOH/LVWRIIRVVLOWD[DSUHVHQWO\FODVVL¿HGDVNeocallichirus (after Schweitzer et al .DUDVDZDDQG1DNDJDZD *DUDVVLQRet al., 2011). Note that N. rodfeldmanni+\åQêLVDUHSODFHPHQWQDPHIRUN. manningi6FKZHLW]HU)HOGPDQQ)DP+HVVLQ+HWULFN1\ERUJDQG5RVV (name preoccupied). Taxon Age Occurrence Remarks N. aetodes Schweitzer et alE (DUO\2OLJRFHQH 3XHUWR5LFR N. agadirensis Garassino et al., 2011 Cenomanian Morocco generic assignment doubtful N. allegranzii Beschin et al., 2005 Middle Eocene Italy generic assignment doubtful N. bona (Imaizumi, 1959) Early–Middle Miocene Japan generic assignment doubtful N. borensis Beschin et al /DWH(RFHQH ,WDO\ N. dijki (Martin, 1883) Late Miocene Phillipines, Java N. fortisi Beschin et al., 2002 Middle Eocene Italy generic assignment doubtful N. hattai.DUDVDZDDQG1DNDJDZD (DUO\±0LGGOH0LRFHQH -DSDQ N. matsoni (Rathbun, 1935) Early Miocene USA (Florida) generic assignment doubtful N. maxima (A. Milne-Edwards, 1870) subfossil Thailand, India (?) N. nishikawai (Karasawa, 1993) Miocene Japan generic assignment doubtful N. okamotoi (Karasawa, 1993) Late Oligocene Japan generic assignment doubtful N. peraensis Collins et al /DWH3OHLVWRFHQH -DPDLFD N. porterensis 5DWKEXQ 2OLJRFHQH 86$ :DVKLQJWRQ2UHJRQ N. quisquellanus Schweitzer et alE 0LGGOH0LRFHQH 'RPLQLFDQ5HSXEOLF JHQHULFDVVLJQPHQWGRXEWIXO N. rhinos Schweitzer and Feldmann, 2002 Middle Eocene USA (California) N. rodfeldmanni+\åQê &DPSDQLDQ &DQDGD %ULWLVK&ROXPELD JHQHULFDVVLJQPHQWGRXEWIXO N. sakiae Karasawa and Fudouji, 2000 Oligocene Japan generic assignment doubtful N. scotti (Brown and Pilsbry, 1913) Oligocene–Pleistocene Caribbean N. wellsi Schweitzer et al /DWH(RFHQH 3DNLVWDQ only on merus, but also on carpus, propodus and dactylus) in the fossil comprehensive study is provided. record for assignment to the genus Neocallichirus. Considering the difficulties with assigning fossil material to extant There are several genera different from Neocallichirus, namely callianassid taxa and different approaches applied when erecting Sergio, Podocallichirus, and Grynaminna, which share very similar fossil taxa, it is fairly possible that the genus Neocallichirus as usually FKHOLSHGV0XFKFRQIXVLRQKDVEHHQFUHDWHGE\6DNDL ZKHQ recognized in the fossil record is a mixture of different genera as the he synonymized Sergio with Neocallichirus and Grynaminna with personal re-examination of some of them by one of the authors (MH) Podocallichirus5HFHQWO\6DNDL UHGH¿QHGDOOWKHVHJHQHUD7KH might show. taxonomic history of all four of them is very complex and until now no general agreement has been achieved. Neocallichirus vs. Sergio Cheliped morphology seen in the genera mentioned above is in palaeontological literature connected virtually with Neocallichirus only. Originally, the genus Sergio was erected to accomodate four species According to De Grave et al. (2009) and Schweitzer et al. (2010) there previously referred to Neocallichirus, based upon the characters of the DUHIRVVLOVSHFLHVLGHQWL¿HGDVNeocallichirus, one fossil species as telson and uropods (Manning and Lemaitre, 1994). The diagnosis was Podocallichirus, one extant species of SergioLVDOVRNQRZQIURPWKH based on four species firstly described by Biffar (1970): Callianassa fossil state and no fossil GrynaminnaLVNQRZQ6LQFHWKHQWZRPRUH trilobata; and Rodrigues (1971): C. guara, C. guassutinga, and C. NeocallichirusVSHFLHVKDYHEHHQGHVFULEHG .DUDVDZDDQG1DNDJDZD mirim. Later, another three species were described under this genus: 2010; Garassino et al., 2011) (see Table 1). Blanco Rambla et al. (1995) introduced Sergio guaiqueri; Manning The morphology of fossil taxa assigned to Neocallichirus is just as and Felder (1995) described Sergio mericeae and reexamined S. variable (and maybe even more) as that of the extant species of the guassutinga; and the last systematic study devoted to the genus Sergio genus. In this context it should be noted that the characteristics of the ZDVWKDWE\/HPDLWUHDQG)HOGHU ZKRGHVFULEHGS. sulfureus. It is chelipeds typical for the genus Neocallichirus were summarized only in worth mentioning that the generic diagnosis was not amended in these several papers dedicated to fossil taxa (Schweitzer and Feldmann, 2002; studies. Sergio was then synonymised with NeocallichirusE\6DNDL Schweitzer et alD VHHDOVR+\åQêDQG+XGiþNRYiVXEPLWWHG KRZHYHUPDQ\ZRUNHUVGLGQRWDFFHSWWKLVV\QRQ\PLVDWLRQ DQGDFFRUGLQJWRSXEOLVKHG¿JXUHVRQO\1RH[KDXVWLYHVWXG\KDVEHHQ (Poore, 2000; Schweitzer et alD 'ZRUVFKDN 3RRUH PDGHWRPDNHDWWHPSWWRHYDOXDWHFKHOLSHGVZLWKLQWKLVVSHFLRVHJHQXV De Grave et al )HOGHUDQG5REOHV 6DNDL According to Schweitzer et al D “assignment of fossils to argued with the presence of intermediate forms between Neocallichirus Neocallichirus must be based on the highly variable morphology of and SergioFRQFHUQLQJWKHQDWXUHRIWHOVRQDQGXURSRGV+HVWDWHG 6DNDL WKHPDMRUFKHOLSHGRIH[WDQWVSHFLHVDQGWUXO\FRQ¿GHQWDVVLJQPHQWRI 2005: 125) that within “the species assigned to Sergio, the character of fossils to the genus will probably only occur when other aspects of the WKHWDLOIDQVKRXOGEHGH¿QHGDVDVSHFL¿FFKDUDFWHUZLWKLQWKHJHQXV body of the animal are recovered”. Many fossil taxa were assigned to Neocallichirus”+H 6DNDL DOVRSRLQWHGRXWRQWKHSUHVHQFH Neocallichirus on the basis of very few characters (mainly on propodus RIWKHPHUDOKRRNLQSergio contrary to the diagnosis of the genus only); thus, revision of all fossil Neocallichirus species referred at some (Manning and Lemaitre, 1994: 40). Schweitzer et al D VXJJHVWHG time to this genus is needed to specify precisely the morphological that palaeontologists should use the presence or absence of a proximal variability of the taxonomically important species and compare it to PHUDOKRRNWRGLVWLQJXLVKEHWZHHQSergio and Neocallichirus. This issue the variability seen in the extant taxa. As in neontological studies the deserves more discussion. genus Neocallichirus is usually recognized on soft part morphology Although Manning and Lemaitre (1994) stated that Sergio does not only, the genus concept as used in palaeontological practice, and thus SRVVHVVDPHUDOKRRN5RGULJXHV LQWKHGHVFULSWLRQRIC. as discussed herein, should be regarded as preliminary one until more guassutinga (type species of Sergio) noted, that“ there are two or three PRGHUDWHO\ORQJVSLQHVIROORZHGE\¿YHVKRUWLUUHJXODUVSLQHVDQGDERXW Table 2. List of extant taxa classified at some time within the genus Sergio Manning and Lemaitre, 1994. eight rounded granules” on the lower margin of merus in males and “WZRODUJHSUR[LPDOVSLQHVIROORZHGE\¿YHWRHLJKWVPDOOHURQHV”in Neocallichirus cacahuate Felder and Manning, 1995 females. Similarly in C. guara,“ the merus has a strong serrated tooth Sergio guaiqueri Blanco Rambla, Liñero Arana and Beltán Lares, 1995 near the proximal extremity of the lower margin wich is denticulated Callianassa (Callichirus) guara Rodrigues, 1971 (Rodrigues, 1971: 211).”Callianassa trilobata and C. mirim are in their Callianassa (Callichirus) guassutinga Rodrigues, 1971 Neocallichirus lemaitrei Manning, 1993 original descriptions characterized also by“ WULDQJXODUYHQWUDONHHORQ Sergio mericeae Manning and Felder, 1995 PHUXV %LIIDU ” in C. trilobata and“ the lower edge produced Callianassa (Callichirus) mirim Rodrigues, 1971 LQWRDODPLQDUVHUUDWHGWRRWK 5RGULJXHV ” on merus in Callichirus monodi de Saint Laurent and Le Loeuff, 1979 C. mirim. Sergio mericeae has major cheliped“ with inferior margin Callianassa pachydactyla A. Milne-Edwards, 1870 of merus armed proximally by elongate process of 2–3 fused spines, Sergio sulfureus/HPDLWUHDQG)HOGHU beyond which is short gap in marginal dentition (Manning and Felder, Callianassa trilobata Biffar, 1970 ” Sergio sulfureus has lower margin of merus“ armed with VPDOOVSLQHVDQGVWURQJEL¿GSURFHVVSUR[LPDOO\ /HPDLWUHDQG)HOGHU ”. reassignment. The only exception is S. guaiqueriZKLFKFOHDUO\ODFNVDQ\SUR[LPDO Reassessment of the genus SergioE\6DNDL LQIDFWLWV PHUDOKRRNVSLQHRUWRRWK %ODQFR5DPEODet al., 1995). However, this ressurection when considering the previous synonymisation) does species was based on only four rather small specimens. Thus, when not really help to resolve the relationship between the taxa discussed assuming that degree of spinosity of chelipeds depends on age of the above, as it is based on variable characters which may change during animal, we can conclude that those specimens represent immature ontogeny, i.e. first two pairs of male pleopods. Already Biffar (1971: stages. This view can be supported with described and figured major PHQWLRQHGWKDWFDOOLDQDVVLGMXYHQLOHVODFNWKHFKDUDFWHULVWLFVKDSH chelipeds of immature specimens of S. sulfureus (Lemaitre and Felder, of the adult appendages. Similarly Felder and Manning (1995) reported ILJVDE DQGS. mericeae (Manning and Felder, 1995: fig. the same for Neocallichirus cacahuate Felder and Manning, 1995 and 5f) which do not have any proximal tooth or spine either, but only a pointed on differences in male pleopod shape between N. cacahuate and VHUUDWHGORZHUPDUJLQ,WVKRXOGDOVREHQRWHGWKDWWKH¿JXUHGPDOHRI N. lemaitrei0DQQLQJ 6DNDL WUHDWVERWKVSHFLHVDVSergio. S. guaiqueriGRHVQRWSRVVHVVVWURQJO\DUPHGGDFW\OXVDQG¿[HG¿QJHU 'ZRUVFKDN DE UHSRUWHGWKHYDULDELOLW\LQWKHVKDSH ZKLFKDUHRWKHUZLVHTXLWHW\SLFDOIRUPDWXUHVSHFLPHQVRIWKHJHQXV RIWKH¿UVWWZRPDOHSOHRSRGVLQN. karumba, N. jousseaumei (Nobili, as described and figured in literature (Rodrigues, 1971; Manning and 1904) (=Callianassa indica de Man, 1905; Neocallichirus taiaro Ngoc- )HOGHU /HPDLWUHDQG)HOGHU ,QFUHDVHGVSLQRVLW\ZLWK Ho, 1995) and N. vigilax GH0DQ N. denticulatus Ngoc-Ho, size (or age) of the specimens was documented also in Neocallichirus 1994), respectively. karumba3RRUHDQG*ULI¿Q Neocallichirus kempi6DNDL The fossil record of the genus Sergio is obscure. Interestingly, Todd E\'ZRUVFKDN 0DMRUFKHOLSHGVRIODUJHUVSHFLPHQVLQWKLV and Collins (2005) recognized Callianassa scotti Brown and Pilsbry, VSHFLHVLQWHUHVWLQJO\SRVVHVVDVLPLODUELRUWUL¿GSUR[LPDOSURMHFWLRQRQ 1913 (=Callianassa vaughani Rathbun, 1918; C. crassimana Rathbun, lower margin of merus. The same conclusion concerning the immaturity 1918; C. miocenica Rathbun, 1919; C. rathbunae Glaessner, 1929) from of the type material of S. guaiqueriZDVUHDFKHGDOVRE\6DNDL the Oligocene to Pleistocene of the Caribbean as a member of the genus 90):“ It seems that the male holotype is immature because the carapace Sergio+RZHYHUXQGHULQÀXHQFHRI6DNDL V V\QRQ\PLVDWLRQRI is only 4.2 mm long, and the male Plp1–2 are undeveloped (Blanco the genus with Neocallichirus, they treated the taxon as Neocallichirus Rambla et al ¿JVDE ” scotti. As such it appeared also in the fossil decapod species list by Finally, recent molecular studies (Tudge et al., 2000; Felder Schweitzer et al. (2010). Another fossil occurrence coming from the and Robles, 2009) resolved Sergio as of paraphyletic nature in the Pliocene–Pleistocene strata of Florida (Portell and Agnew, 2004) has arrangement presented above (see also Table 3). In Felder and Robles EHHQLGHQWL¿HGDVH[WDQWSergio trilobatus (Biffar, 1970). (2009: fig. 1) two tested taxa, S. mericeae and S. trilobata were As shown above, the problem of distinguishing Neocallichirus and positioned in distinctly different branches. Sergio mericeae (considered Sergio from one another in the fossil record is more-less a matter of to be a close relative of the type species of Sergio) was allied with their definition as biologically defined taxa. In this respect, the fossil otherwise monophyletic Neocallichirus, S. trilobata was, however, record can tell only little to solve this issue. positioned basally to the grouping of genera Corallianassa Manning, 1987, Glypturus6WLPSVRQGrynaminna and Neocallichirus. Neocallichirus vs. Podocallichirus Felder and Robles (2009: 334) noted that such arrangement“ raises a TXHVWLRQDVWRWKHYDOLGLW\RIWKHJHQXVDQGUHJDUGOHVVRIWKDWLVVXH The present status of the genus Podocallichirus is rather confusing. argues for generic reassignment of S. trilobata”6DNDL HUHFWHG ,WZDVHUHFWHGE\6DNDL WRDFFRPRGDWHVHYHQVSHFLHVSUHYLRXVO\ a new genus, Glypturoides, for this taxon. We agree with this generic mostly treated as Callichirus sensu Le Loeuf and Intès (1974) (see Table 3. Taxonomic history of generic assignment of extant taxa treated as Sergio Manning and Lemaitre, 1994. Taxon Sakai (1999) Tudge et al. (2000) Sakai (2005) Poore (2010) Sakai (2011) N. cacahuate Neocallichirus Neocallichirus Neocallichirus Neocallichirus Sergio S. guaiqueri Neocallichirus Sergio Neocallichirus Sergio Neocallichirus C. guara Neocallichirus Sergio Neocallichirus Sergio Neocallichirus C. guassutinga Neocallichirus Sergio Neocallichirus Sergio Sergio N. lemaitrei Neocallichirus Neocallichirus Neocallichirus Neocallichirus Sergio S. mericeae Neocallichirus Sergio Neocallichirus Sergio Sergio C. mirim Neocallichirus Sergio Neocallichirus Sergio Neocallichirus C. monodi Neocallichirus Callichirus Neocallichirus Neocallichirus Sergio C. pachydactyla Neocallichirus Neocallichirus Neocallichirus Neocallichirus Sergio S. sulfureus Neocallichirus Sergio Neocallichirus Sergio Sergio C. trilobata Neocallichirus Sergio Neocallichirus Sergio Glypturoides Tables 4 and 5). Podocallichirus as understood in the classification as proximally positioned serrated meral blade, however, in general, of De Grave et al. (2009) apparently comprises these seven taxa. the states of these characters overlap in many cases with typical Poore (2000) erected the genus Grynaminna to accomodate the new Neocallichirus species. Contrary to that, Podocallichirus sensu stricto species G. tamakii. This genus was considered as a junior synonym of comprises very unusual species, whose minor cheliped and also some PodocallichirusE\6DNDL /DWHUKH 6DNDL UHFRJQL]HG characters of the major one, is different than majority of callianassid Grynaminna as a separate genus and divided PodocallichirusLQ¿YH WD[D VHH/HQ]DQG5LFKWHUV ¿JV YLUWXDOO\WKHRQO\VSHFLHV independent genera; Podocallichirus remained monotypic containing with broadly similar minor chela has been described as Lepidophthalmus only its type species (Callianassa madagassa Lenz and Richters, 1881). socotrensis6DNDLDQG$SHO 1RWKLQJVLPLODUKDVEHHQIRXQG The remaining species are accomodated within the newly established LQWKHIRVVLOUHFRUGVRIDU$VDFRQVHTXHQFHZKHQGHDOLQJZLWK genera Balsscallichirus, Barnardcallichirus, Forestcallichirus and fragmentary material, Podocallichirus sensu latoZRXOGEHLGHQWL¿DEOH Tirmizicallichirus. Their recognition as separate genera is obscure, as LQWKHIRVVLOUHFRUGDVDPHPEHURIEURDGO\GH¿QHGNeocallichirus. they are based on characters changing during ontogeny (see above), and thus conclusions based on their nature can be misleading. It is worth Neocallichirus vs. Grynaminna mentioning that restriction of Podocallichirus to C. madagassa only is VXSSRUWHGDOVRE\3RRUH +HFODVVL¿HVWKHUHVWRIPodocallichirus The monotypic genus Grynaminna was erected for a distinct species species sensu6DNDL DVPHPEHUVRIEURDGO\GH¿QHGCallichirus from Kyushu, Japan. As defined by Poore (2000) it is very similar to (Table 5). Neocallichirus, however, it differs in the nature of antennae, uropods $VDUHVXOWZHFDQWDONDERXWWZRGLVWLQFWO\GLIIHUHQWFRQFHSWVRI DQGSOHRSRGV 3RRUH ,WODFNVWKHPHUDOKRRNRQPDMRU the genus Podocallichirus RULJLQDOFRQFHSWJLYHQE\6DNDL FKHOLSHGDQGSRVVHVVHVQRQWDSHULQJFKHODHZKLFKDUHRWKHUZLVHTXLWH comprising type species plus six other species, here referred as typical for Neocallichirus and Sergio. Podocallichirus sensu lato; and 2) concept of the monotypic genus It seems that chelipeds of Grynaminna tamakii Poore, 2000 (type Podocallichirus comprising the type species, Callianassa madagassa, species of Grynaminna) are sexually dimorphic. In males there is a only, thus Podocallichirus sensu stricto (supported by Poore, 2010 and QRWFKDWWKHEDVHRIWKH¿[HG¿QJHURIPDMRUFKHODZKLFKLVQRWSUHVHQW 6DNDL :KHQFRPSDULQJFKHOLSHGVRIWKHVHWZRFRQFHSWVRQH in females. In males the dactylus is armed with several teeth, whereas in FDQKDYHGLI¿FXOWLHVWR¿QGDVXLWHRIFRPPRQFKDUDFWHUVZKLFKDUHQRW IHPDOHVLWLVVWUDLJKW 3RRUH ¿J present also in Neocallichirus. Podocallichirus sensu lato possesses 6DNDL V\QRQ\PLVHGGrynaminna with Podocallichirus6DNDL several characters which are not very common in Neocallichirus, such KRZHYHULQKLVODWHUZRUN 6DNDL KHUHFRJQL]HGLWDVD distinct genus (see Table 5). We concur. Table 4. List of extant taxa classified at some time within the genus The fossil record of Grynaminna is obscure, however, we argue that Podocallichirus6DNDL “Neocallichirus”grandis.DUDVDZDDQG*RGD IURPWKH0LGGOH Callianassa (Callichirus) balssi Monod, 1933 Pleistocene of Japan (Aichi Prefecture) can be accomodated within the Callichirus foresti Le Loeuff and Intès, 1974 genus. Although very well preserved, the material comprises chelipeds Callianassa gilchristi Barnard, 1947 only. Based on cheliped morphology Obata and Hayashi (2001) removed Callianassa (Callichirus) guineensis de Man, 1928 “N.” grandis to Grynaminna. Later, Karasawa et al IROORZLQJ Callianassa madagassa Lenz and Richters, 1881 WKHZRUNRI6DNDL UHPRYHGWKHVSHFLHVWRPodocallichirus and Callianassa (Callichirus) masoomi Tirmizi, 1970 as such the species appeared in a list of fossil decapod crustaceans by Grynaminna tamakii Poore, 2000 Callichirus tenuimanus de Saint Laurent and Le Leouff, 1979 Schweitzer et al 1RZDIWHUWKHSXEOLFDWLRQRI6DNDL V systematic reconsideration of callianassoid ghost shrimps one has GLI¿FXOWLHVWRMXGJHKRZWRFODVVLI\³Neocallichirus” grandis. Virtually Table 5. Taxonomic history of generic assignment of extant taxa treated as Podocallichirus6DNDL Taxon Sakai (1999) Tudge et al. (2000) Sakai (2005) Poore (2010) Sakai (2011) C. balsii Podocallichirus Callichirus Podocallichirus Callichirus Balsscallichirus C. foresti Podocallichirus Callichirus Podocallichirus Callichirus Forestcallichirus C. gilchristi Podocallichirus Callianassa Podocallichirus Callichirus Barnardcallichirus C. guineensis Podocallichirus Callichirus Podocallichirus Callichirus Balsscallichirus C. madagassa Podocallichirus Callianassa Podocallichirus Podocallichirus Podocallichirus C. masoomi Podocallichirus Callianassa Podocallichirus Callichirus Tirmizicallichirus G. tamakii - Grynaminna Podocallichirus Grynaminna Grynaminna C. tenuimanus Podocallichirus Callichirus Podocallichirus Callichirus Barnardcallichirus WZRRSWLRQVDUHDYDLODEOH LIIROORZLQJ3RRUH DQG6DNDL as undistinguishable when dealing with chelipeds only. In that case in recognition of Grynaminna as a distinct genus, and considering it should be classified within the genus Neocallichirus (as originally the similarity between N. grandis and G. tamakii as of taxonomic proposed) in the broadest sense, thus as defined by Manning and importance, then “N.” grandis should be classified within the genus )HOGHU :HDVVXPHWKDWDWWKHSUHVHQWVWDWHRINQRZOHGJHWKH Grynaminna as recognized by Obata and Hayashi (2001; 2) another FODVVL¿FDWLRQRI³N.”grandis within the genus Podocallichirus seems option is to consider Neocallichirus, Grynaminna and Podocallichirus to be misleading. Fig. 1. Grynaminna grandis .DUDVDZDDQG*RGD IURPWKH0LGGOH3OHLVWRFHQH$WVXPL*URXS -DSDQ $±%ERWKFKHOLSHGVRIDSUHVXPHGPDOH specimen in lateral (A) and dorsal view (B) (MFM142381), note serrated lower margin of propodus (B), which is preserved turned over in contrast to the rest of the cheliped; C–D, both chelipeds of a presumed female specimen in mesial (major chela) and lateral view (minor chela) (MFM142497), QRWHNHHOHGXSSHUDQGORZHUPDUJLQVRISURSRGXVDQGFDUSXV (LVFKLXPPHUXVDQGFDUSXVIURPPHVLDOYLHZ 0)0 QRWHVHUUDWHGORZHU margin of the merus; F, remains of both chelipeds of the same specimen (MFM142500); G, both chelipeds of a presumed male specimen (MFM142499), major cheliped in lateral view and minor cheliped in mesial view; H, major cheliped of a presumed female specimen in mesial view (MFM142498). All specimens were covered with ammonium chloride prior the photography. All specimens are preserved within the burrows. specimens supposedly attributed to females do not possess a distinct Systematic palaeontology QRWFKDQGWKHDUPDWXUHRI¿QJHUVLVQRWZHOOSURQRXQFHG )LJV&+ It seems that the genus Grynaminna is biogeographically restricted Order Decapoda Latreille, 1802 WR-DSDQ 3RRUH 6DNDL 6RIDUWKHUHLVRQO\RQHUHSRUWHG Infraorder Axiidea de Saint Laurent, 1979 occurrence of extant NeocallichirusIURP-DSDQ6DNDL Family Callianassidae Dana, 1852 2011) reported N. indicus (=N. jousseaumei IURP WKH 5\XN\XV Subfamily Callichirinae Manning and Felder, 1991 'ZRUVFKDN D GLGQRWOLVWWKLVRFFXUUHQFHEHFDXVHRILWV XQUHVROYHGVSHFLHVLGHQWLW\ 3&'ZRUVFKDNSHUVFRPP 7KH Genus Grynaminna Poore, 2000 fossil record of the genus NeocallichirusLQ-DSDQH[WHQGVEDFNWRWKH Type species: Grynaminna tamakii Poore, 2000, by original 0LRFHQH .DUDVDZDDQG1DNDJDZD 7KHH[WDQWDQGH[WLQFW designation and monotypy. Podocallichirus sensu lato and Sergio has not yet been reported from Included fossil species: Grynaminna grandis (Karasawa and Goda, Japan. Diagnosis: See Poore (2000: 150). We argue that the genus concept as Conclusions SUHVHQWHGE\6DNDL LHEDVHGRQWKH¿UVWWZRSDLUVRIPDOH pleopods is not enable (see above). From the present contribution several conclusions can be made: 1) Generic assignment of fossil callianassid remains can be done Grynaminna grandis (Karasawa and Goda, 1996) successfully on the basis of chelipeds only, but only if all cheliped (Fig. 1A–H) elements (ischium, merus, carpus, propodus and dactylus) are at hand. CallianassaVS.DWRDQG.RL]XPL S ¿J± Thorough comparisons between fossils and extant material should be Calliax VS.DUDVDZDDQG7DQDND S¿J± always made before assigning the fossil remains to the genus level. Neocallichirus VS.DUDVDZD1RKDUDDQG6KLPRML S¿J 2) More comparative studies among the extant callianassids using “Neocallichirus” grandis.DUDVDZDDQG*RGD S¿J .DUDVDZD the chelipeds is needed to recognize important characters which are SSO¿JV± UH¿JXUHGIURP.DUDVDZDDQG*RGD Neocallichirus grandis Karasawa and Goda; Kato and Karasawa, 1998, p. 5, consistent troughout all the members of the genus. In this respect a SO¿JV± 6FKZHLW]HUet alES LQDOLVW SDSHUE\6DNDL RQWKHPDMRUFKHOLSHGYDULDWLRQVLQVRPH-DSDQHVH Grynaminna grandis .DUDVDZDDQG*RGD 2EDWDDQG+D\DVKLS callianassids is very useful for palaeontological practice. Manning and ¿J± Felder (1991) pointed out that some characters occurring on the merus Podocallichirus grandis (Karasawa and Goda); Karasawa et alS (usually in the combination of other features of hard part morphology) SO¿JV± 6FKZHLW]HUet al., 2010 p. 39 (in a list). can be used for generic assignment; since then, however, nearly Material examined: MFM142381, MFM142497–142500 deposited at WZHQW\ QHZH[WDQWFDOOLDQDVVLGJHQHUDZHUHGHVLJQDWHGLQPDQ\FDVHV the Mizunami Fossil Museum, Gifu, Japan. without any or little attention paid to the nature of the chelipeds. Remarks: The morphology of major and minor chelipeds of 3) Revision of all fossil species referred to Neocallichirus is needed “Neocallichirus” grandisKDVEHHQVXI¿FLHQWO\GHVFULEHGE\.DUDVDZD to specify precisely the morphological variability of taxonomically DQG*RGD 7KHVSHFLHVKDVPDMRUPRUSKRORJLFDOVLPLODULWLHV important characters and compare it to the variability seen in extant to Grynaminna tamakii Poore, 2000, as was already pointed out by Neocallichirus species. Obata and Hayashi (2001). The similarities include characters on the major cheliped: the ovoid-shaped merus with serrated lower margin Acknowledgements DQGZLWKRXWDSURQRXQFHGPHUDOEODGHRUKRRN QHDUO\TXDGUDWHFDUSXV short, triangular fixed finger; and the dactylus armature in supposed :HZRXOGOLNHWRH[SUHVVWKHGHHSHVWWKDQNWR3HWHU&'ZRUVFKDN male specimens (see below). Grynaminna grandis bears a large, blunt (Natural History Museum, Wien, Austria) for fruitful discussions triangular tooth on the fixed finger, which G. tamakiiODFNV,QWKLV on systematics and biology of the extant Callianassidae and helpful FDVHWKHFKDUDFWHUVRQWKH¿[HG¿QJHUFDQEHFRQVLGHUHGRIWD[RQRPLF FRPPHQWVRIWKHHDUOLHUYHUVLRQRIWKHPDQXVFULSW7KHZRUNKDVEHHQ importance on the species level. Otherwise, the overall cheliped VXSSRUWHGE\UHVHDUFKJUDQWV$399WR'DQLHOD5HKiNRYi dimensions and ratios are virtually the same in both species. Thus, &RPHQLXV8QLYHUVLW\%UDWLVODYD6ORYDNLD .(*$. WR “Neocallichirus” grandis can be best accomodated within the genus 1DWiOLD+XGiþNRYi &RPHQLXV8QLYHUVLW\%UDWLVODYD6ORYDNLD DQG Grynaminna. &RPHQLXV8QLYHUVLW\*UDQW8.DQG3DO6,536HSNRVNL*UDQW The material of Grynaminna grandis comprises two morphotypes WR0DW~ã+\åQê which seem to mirror sexual dimorphism and can be compared with sexually dimorphic chelipeds of G. tamakii (i.e., Kato and Karasawa, References 2009; Kato, in preparation). The major propodus of G. grandis supposedly attributed to male has a distinct notch at the base of the Barnard, K. H. (1947), Descriptions of new species of South African decapod ¿[HG¿QJHUDQGDUPHGERWK¿[HG¿QJHUDQGGDFW\OXV )LJ* ZKHUHDV Crustacea, with notes on synonymy and new records. The Annals and 0DJD]LQHRI1DWXUDO+LVWRU\VHULHV>IRU @± in the Estuarine Ghost Shrimp Callianassa louisianensis Schmitt, Beschin, C., A. Busulini, A. De Angeli, and G. Tessier (2002), Aggiornamento 1935. 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