JOURNAL OF CRTJS I AC E-\N UIOLOGY. l3(3): ,143-,155. 1993 GEOLOGICAL HISTORY OF BRACHYURAN DECAPODS FROM NEW ZEALAND Rodnev M. Feldmann and Colin L. McLav ABSTRACT Recently completed, comprchensive surveys of the living and fossil brachyurans from New Zealand pcrmit an analysis ol thc palcobiogeographic hisrory of the group. Among the l8 families known from the modern launa, 500/o arc rcpresented in the lossil record. The modern launa is con.rprisedof lewer family-lcvcl taxa lhan are present in Japan, China, or Wesl Africa, which rellects limitc'd low latitude influence.Development of some elemenlsof the New Zealand brachyuran fauna can be lraced into thc (lrctaceous. As early as thc Eocene, a characteristic fauna, with high latitudc, lemperate affinities, had dcveloped. Teth1,an, or tropical and sub- tropical. taxa are less common than previously presumed. The New Zealand brachyuran fauna was eslablishcd largely prior 10 separation ofthe southern conlinents and development ollhc circum-Antarctic current svstem. The living and fossil crab launa of New Zealand has a distinctivc character which has resulted from early Tertiary appearance of families from nearby southern hemi- spherc regions and subsequent develop- rnenl in relativcl)i grealcr isolalion. The lau- na ls moderately well known. Since the lirst living species was described, lVototnithra.x ,1r.fr/-r (Herbst, 1788),a voluminous litera- ture has dealt with the description of ncw speciesand characterization of certain as- pects of the fauna. Similarly, subsequcntto the description of the hrst fossil crab from New Zealand. Ttuttidocarcinus tumidtts (Woodward, 1876). more than 38 papers havc referred to lossil crabs from thc region. Recently, data regarding the recent fauna have been compiled (Mcl-ay, 1988) and all known references to fossils havc been brought together (Feldmann and Keyes, 1992). In addition, all fossil material de- posited in university and public collections has been examined and identificd. Thus. for thc first time, a large proportion of the lit- erature dealing with Ncw Zealand crabs has been examined, reinterpreted, and com- piled. The origin and subsequent history of this fauna has been of considerable interest to biogeographers, owing to the complex his- tory of arrival of taxa to New Zealand. Dell (1968) provided the lirst detailed analysis of the distribution and biogeographicaffin- ities of Brachyura around New Zealand. Fleming (1962, 1979) postulared rhar rhe "typical" New Zealand marine decapod faunas appearcd subsequent to the Oligo- cene. This conclusion was based upon the fossil record, as it was then understood (Glaessner,I 960). In the conte xt of Glaess- ner's pioneering study of New Zealand decapods,Fleming concluded thal the affin- ities of the decapods were primarily with warm watcr. Australian or Malayo-Pacific (Tethyan), forms. Newman (1991) has re- ccntly reinlorccd this conclusion by sug- gesting that many New Zealand decapod taxa may be relicts of a former amphitrop- ical distribution. This pattern came into question when Fcldmann ( I 984) describedthc crablike an- onruran Haumuriaegla glaassneri from ma- rine Cretaceousrocks in New Zcaland and concluded that this spccieswas anccstral to the living aeglids which are exclusively in- habitants of fresh-water habitats in south- ern South America. This history empha- sized the role of the southern high latitudes as a sourceofnew decapodtaxa. Subsequent field work, primarily in Eocene and Creta- ceous rocks of New Zealand, coupled with sludies on fossil decapods from Antarctica, now suggest that this pattern may bc wide- spreadand that Fleming's conclusions musl be modified. Thus, the purpose of this work is to examine the relationships between the living and fossil crabs, the brachyuran dec- apods, of New Zealand to reevaluate the paleobiogeographic history of the fauna. THe BRAcHyURANRpcoRo The Extanl Brachyuran Fauna The modern decapod fauna of New Zea- land and surrounding waters includes at least 443
Transcript
J O U R N A L O F C R T J S I A C E - \ N U I O L O G Y . l 3 ( 3 ) : , 1 4 3 - , 1 5 5 . 1 9 9 3
GEOLOGICAL HISTORY OF BRACHYURAN DECAPODSFROM NEW ZEALAND
Rodnev M. Feldmann and Colin L. McLav
A B S T R A C T
Recently completed, comprchensive surveys of the living and fossil brachyurans from NewZealand pcrmit an analysis ol thc palcobiogeographic hisrory of the group. Among the l8families known from the modern launa, 500/o arc rcpresented in the lossil record. The modernlauna is con.rprised of lewer family-lcvcl taxa lhan are present in Japan, China, or Wesl Africa,which re l lects l imi tc 'd low lat i tude inf luence. Development of some elemenls of the New Zealandbrachyuran fauna can be lraced into thc (lrctaceous. As early as thc Eocene, a characteristicfauna, with high latitudc, lemperate affinities, had dcveloped. Teth1,an, or tropical and sub-tropical. taxa are less common than previously presumed. The New Zealand brachyuran faunawas eslabl ishcd largely pr ior 10 separat ion of the southern conl inents and development o l lhccircum-Antarctic current svstem.
The living and fossil crab launa of NewZealand has a distinctivc character whichhas resulted from early Tertiary appearanceof families from nearby southern hemi-spherc regions and subsequent develop-rnenl in relativcl)i grealcr isolalion. The lau-na ls moderately well known. Since the l irstl iving species was described, lVototnithra.x,1r . f r / - r (Herbst , 1788), a voluminous l i tera-ture has dealt with the description of ncwspecies and characterization of certain as-pects of the fauna. Similarly, subsequcnt tothe description of the hrst fossil crab fromNew Zealand. Ttut t idocarc inus tumidt ts(Woodward, 1876). more than 38 papershavc referred to lossil crabs from thc region.Recently, data regarding the recent faunahave been compiled (Mcl-ay, 1988) and allknown references to foss i ls havc beenbrought together (Feldmann and Keyes,1992). In addition, all fossil material de-posited in university and public collectionshas been examined and identif icd. Thus. forthc first t ime, a large proportion of the l it-erature dealing with Ncw Zealand crabs hasbeen examined, reinterpreted, and com-pi led.
The origin and subsequent history of thisfauna has been of considerable interest tobiogeographers, owing to the complex his-tory of arrival of taxa to New Zealand. Dell(1968) provided the l irst detailed analysisof the distribution and biogeographic affin-it ies of Brachyura around New Zealand.Fleming (1962, 1979) postu lared rhar rhe"typical" New Zealand marine decapodfaunas appearcd subsequent to the Oligo-
cene. This conclusion was based upon thefossil record, as it was then understood(Glaessner, I 960). In the conte xt of Glaess-ner's pioneering study of New Zealanddecapods, Fleming concluded thal the affin-it ies of the decapods were primarily withwarm watcr. Australian or Malayo-Pacific(Tethyan), forms. Newman (1991) has re-ccntly reinlorccd this conclusion by sug-gesting that many New Zealand decapodtaxa may be relicts of a former amphitrop-ical distribution.
This pattern came into question whenFcldmann ( I 984) described thc crablike an-onruran Haumuriaegla glaassneri from ma-rine Cretaceous rocks in New Zcaland andconcluded that this spccies was anccstral tothe l iving aeglids which are exclusively in-habitants of fresh-water habitats in south-ern South America. This history empha-sized the role of the southern high latitudesas a source ofnew decapod taxa. Subsequentfield work, primarily in Eocene and Creta-ceous rocks of New Zealand, coupled withsludies on fossil decapods from Antarctica,now suggest that this pattern may bc wide-spread and that Fleming's conclusions muslbe modified. Thus, the purpose of this workis to examine the relationships between theliving and fossil crabs, the brachyuran dec-apods, of New Zealand to reevaluate thepaleobiogeographic history of the fauna.
THe BRAcHyURAN RpcoRo
The Extanl Brachyuran Fauna
The modern decapod fauna of New Zea-land and surrounding waters includes at least
443
444 J O I . J R N A L O F C R I J S T A ( - E A N B I O I - O ( ; Y . V O L . I 3 . N O . 3 . I 9 9 3
Table l. Systematic list of the genera of crabs known from New Zealand and the families of brachyurandccapods. Occurrences oftaxa known from the fossil record are indicated by an asterisk. The geographic localities,following the lamily names, indicale the presence of thosc lamilies in New Tnaland, China (Dai and Yang,199 l), Japan (Sakai, 1976), and Chile (Retamal, 1981; B6ez and Martin, 1989). New Zealand occurrences aretaken from the literature cited throughoul the paper. The occurrence of Chaceon (Geryonidae) is from Manningct a l . (1990). The l is t of exlant fami l ies was laken l rom Manning and Hol thuis (1981) and Guinot (1991).
Infraorder BrachyuraSection Podotremata
Family Homolodromiidae-Ncw Zealand, Japan, ChileHomolodromia*
Family Dromiidae-New Zealand, Japan, China, ChileDromia
Family Dynomenidae- JapanFami ly Homol idae-New Zealand, Japan, Chi le
HomolaParamola
Family PoupiniidaeFam ily Latreilli idae - New Zealand, Japan, China
LatreilliaFamily Raninidae-New Zealand, Japan, China
Lyreidus, Lyreidus*Laeyiranina*Hemioon*
Fami ly Cymonomidae-New Zcaland, ChinaCvmonomus
Family Torynommidae- New ZealandTorvnomma*EodorippC
EurynomeCyrtornaiaPlatynniaPyromaiaAchaeopsisAcheusRochiniaEurynolambrusLeptornaiaTrichoplatusTeratornaiaThacanophrvsNot o,n i t hrax, N ol ct mit hrax*Lept ornit h rar, Lept o m it hrax*J acqui not ia, Jacqui not ia*,{ct inolo(arci nus*?Micrttmithrax*
Family MimilambridaeFamily Parthenopidae-Japan, ChinaFami ly Bel l i idae-New Zealand, Chi le
HeteroziusFami ly Leucosi idae-Ncw Zealand, Japan, China, Chi le
EbaliaMerott'ltplusRandallia
Section ThoracotremataFami ly Gecarcin idae- Japan, ChinaFamily Grapsidae - N ew Zealand, .lapan, Ch i na, Chi le
78 species of brachyurans. arrayed in 53genera and l8 families, including the La-treil l i idae as distinct from the Homolidae(see Mcl-ay, 1988) (Table I ). Subsequenl to
that work. several other crab taxa have beencollected from New Zealand waters, butthere has been no published documentationof them. Although these new discoveries wil l
446 JOLJRNAL OF ( RUSTAC'EAN BIoLOGY. VOL I . ] . N( ) . 3 . I993
increase the number of species and, prob-ably, genera, there wil l be l itt le or no changein the number of families, except those thatarise from reconsideration and elevation ofsubfamilies. The faunal diversity as deter-mined at the family level is moderate. Man-ning and Hol thuis (1981) recognized 36families of l iving brachyurans and Bowmanand Abele (1982) recognized 47 families,the sole difference between the two lists be-ing in the primarily fresh-water superfamilyPotamoidea. Subsequent ly , Guinol (1991)named the Poupiniidae. Thus, the New Zea-land fauna includes 49o/o of the 37 extantfamilies, using the family l ist of Manningand Holthuis or 38o/o of the extant families,using the family l ist of Bowman and Abele.
The number of genera and species is rel-atively low, probably reflecting the cooltemperate water conditions and the relativegeographic isolation. Although it is diff icultto find comparable areas with which corn-parisons can be made, and relatively few ofthese useful, comprehensive surveys havebeen conducted in certain areas and thesecan serve as a basis lor comparison. Reta-mal (1981) surveyed the decapod faunasknown from Chile and recorded 130 speciesin 70 genera and l6 families. He includedBellia Milne Edwards, 1848, in the Atele-cyclidae, rather than in the Bell i idae. Sub-sequent 1o this work, B6ez and Martin ( 1989)added anotlrer species, within a previouslyunreported family, the Homolodromiidae.bringing the total number of families to 18.In thc Gal6pagos Is lands, Garth (1991) not-ed I 20 species in 9 I genera. The brachyuranfauna of the easlern United States was sum-mar ized by Wi l l iams (1984) who docu-mented 174 species in 100 genera and 18families. In these surveys the number ofgenera and species far exceeds the numbersin New Zealand, although the number offamilies is similar.
A recent compilation of the crabs alongthe coast of China (Dai and Yang, 1991)reported 604 species ofcrabs 1n 226 generaand 26 fami l ies. Sakai (1976) l is ted ap-proximately 900 species in 328 genera ofcrabs in 3l families in Japan, and Manningand Hol thuis (1981) l is ted 218 species in120 genera and 26 families along the westcoast of Africa. In all. l2 families of brachy-urans occur in Japan or China which are notrecorded from New Zealand. Ctne family.
the Belliidae. is known from New Zealandand Chile but not from Japan or China. Theremaining lamilies are known from bothNew Zealand and either China or Japan.
Much of this variation undoubtedly canbe attributed to differences in area surveyedand variations in taxonomic concepl. Noattempt has been made 1o assess these vari-ables. Two factors which clearly affect thecomparisons are isolation and geographicposition. West Africa, China, Japan, andChile are all situated in close proximity toother faunal regions, as they have beenthroughout the Cenozoic, so that dispersalofnew taxa into those areas could have oc-curred with ease. By contrast, New Zealandhas been relatively isolated from other landmasses since the Oligocene (Smith e/ a/.,1981), reducing the probabil ity of infusionof new stocks. Furthermore, New Zealandlies entirely within temperate and cool-tem-perate waters, but the other areas have trop-ical and subtropical. as well as temperate,influences.
The species composition of the New Zea-land fauna is also dislinctive. Forty-six spe-cies, 590/0, of the fauna are in three families,the Majidae (22 species. 28olo), the Hyme-nosomatidae (14 species, 180/o), and theGrapsidae (10 species, 130/o). Although theMajidae and Grapsidae tend to be well rep-resented in nearly all crab faunas, the Hy-n'lenosomalidae are unusually abundanl inNew Zealand and thc Xanthidae. Leucosi-idae, and Calappidae, for example, are ex-tremely restricted. The xanthids and leu-cosiids account for only 9o/o of the species.Tlrere are no published records ofcalappids,although there is one collection taken fromNew Zealand waters and, as yet, unpub-lished. The Bell i idae are represented by onlya single species, lol0, in New Zealand. Thisfamily is represented jn Chile by a singlespecies (Retamel , 198 I ) , but is unknown inWest Africa, China, and Japan.
Endemic i t y
In the l iving brachyuran fauna, none ofthe 18 famiiies is endemic. At the genericlevel, 9 genera (l7o/o) are endemic and 39species (500/o) are not known outside NewZealand walers. All of these species live onthe continental shelf or in shallow coastalwaters. None of the deep-water species isendemic. Because the conceDt of subfamilv
F E L D I \ I . A , N N A N D M c L A Y : C ; E O L O a i l a A L l i I S T L ) R Y O F N E w Z E A L A N D I I R A C H Y L T R , q N S 4 4 1
has not been evenly applied to all groups,it is not convenient to make comparisonsat that level in all families. However, amongthe Majidae, none of the subfamilies rep-resented in the fauna is endemic.
The endemic genera include Eurltnolam-b rus M i l ne Edwards and Lucas , 1843 ,Trichoplatus Milne Edwards, 1876, Tera-tomaia Griff in and Tranter, 1986, and Jac-quinot ia Rathbun. 1915, in the Maj idae.Pteropeltarion Dell. 1972. in the Atelecy-clidae, Helerozius Milne Edwards. 1867. inthe Bell i idae. l, ' teommalocarcinus Takedaand Miyake. 1969, in the Goneplacidac. andHalimena Melrose, 1975, and Neohymen-icus Lucas, 1980, in the Hymenosomatidae .The endemic species belong mostly to theMajidae (l l) and the Hymenosomatidae(10). Four of the majid genera (26.70/o) andtwo of the hymenosomatids (33.30/o) are en-demic. This h igher level of endemic i tyamong the hymenosomat ids is a lso ev idenlat the species level ; 71.4olo occur only inNew Zealand whereas 500/o of the majidsare known only from the area. The relativelylarge number of species within these fami-l ies may have resulted from population iso-lation and localized current patterns result-ing f rom the changes in land-massconhguration during the Tertiary (Fleming.1979).
The nearest large land mass to New Zea-land is Australia and 94o/o of families, 770loof genera, and 4oo/o of New Zealand speciesalso occur across the Tasman (Mcl-ay. 1988).The New Zealand genera which are abscntfrom Australia arc Pyromaia Stimpson,187 1, Eurynolambrus, Leptontaia Griflinand Tranter, 1986, Trichoplatus, Terato-ntaia, Jacquinotia in the Majidae, Ptero-peltarion in the Atelecyclidae, Cancer Lrn-naeus, 1758, in the Cancridac, Heteroziusin lhe Bell i idae, lt lettmmqtocarinus in theGoneplacidae, and Halirnena and Neoht'-menicus within the Hymenosomatidae. Thegenus Calccr is included here, althoughNat ions (1975) recorded i ts presence in theregion of Tasmania and southcastern Aus-tralia. In fact. the genus probably did notoccur in Australia prior to man's introduc-ing of it to Tasmania early in this century(McNeill and Ward, 1930). These generamay have evolved in, or been introducedinto, New Zealand sometime after its sep-aration fronr Australia. However. the fossil
record does not provide adequate resolulionto test this hypothesis at this time. BothHeterozius and Cetnt'er must have historicalties with South America because many oth-er species in their respective families occurin that region.
A high level of endemicity is not foundin the Grapsidae, where there are no en-demic genera and only 2 endemic species,Hemigrapsus edwardsi (Hilgendorf, 1882)and Helice crassa Dana. 1851. Some haveco lon i zed Sou th Amer i ca , f o r examp leHemigrapsus t 'renLtlatr.rs (H. Milne Ed-wards, 1837) and C),clograpsus lavauri H.Milne Edwards, 1853. Grapsids typicallyhave rather extended larval periods whichmay explain their widespread distributionand low endemicity. A similar story is foundamong the Portunidae which have largenumbers of larval stages. Within this familythere are only two endemic species, .Ay'ec-lo(arcinus antarct icus (Jacquinot, I 853) andN. bennet t iTakeda and Miyake, 1969. bothfrom cold waters south of New Zealand.
The Xanthidae, an undoubted tropica^group, are poorly represented in the faunawith only four species, only two of whichare endemic, Pilumnus lumpinus Bennett,\964, and P. novaezelandiae Fllhol. 1886.There are no apparenl descendants of themost common and best known fossil xan-thid, T-umidocarcinus Glaessner, 1960, inthe launa today. Thejr ext inc l ion may berelated to paleoceanographic changes relat-cd to general cooling following the Mioceneand lcading to the subsequent periods ofglaciation.
Finally, Halicarcinus White, 1846, Cv-clograpsus Milne Edwards, 1834, Lepto-grapsus Milne Edwards, 1853, PlagusiaLa-treil le, 1804, Hemigrctpsus Dana, 185 l, andOvalipes Rathbun, 1898. exhibit circum-polar distribution. Unfortunately, there isno fossil record available in New Zealandto trace the history of this pattern.
The Fossil Brachyuran Fauna
At least 39 species of brachyurans, in 26genera and l2 families, have been describedfrom the fossil record (Feldmann and Keyes,1992) (Table 2). Of the families, nine (500/o)are known from both the fossil and recenlrecord and three, the Homolodromiidae,Torynommidae, and Calappidae, are not re-corded as having l iving descendants in New
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sils. Thus, although there is no statisticalbasis for comparison, the number of con-generic fossil and Recent records from NewZealand would seem to suggesl rather closecorrespondence.
A good correspondence can also be sug-gested at the specics level within some fam-llies. J ac q ui not i a edv, a rd si (Jacq ui not, I 8 5 3 ).Notomithrax minor (F|lhol, I 885), and Lep-tomithrax sp., in the Majidae, Macrophthal-tnott,t hirt ipes (Heller, 1862) in the Ocypod-idae, Cancer norat,ze landiae (Jacquinot,1853) in the Cancridae, Ovalipes punctatus(de Haan, 1833) in the Portunidae, and pos-slbly Carcinoplax sp.. in the Goneplacidae,have all been reported from Pleistocene oc-currences (Feldmann and Keyes, 1992). Thegrapsids are notably underrepresented asonly a single occurrence of the famlly, Mio-grapsus pqpaka Fleming, 1981, from theMiocene. is known.
Geological investigations within the pastdecade have been focused primarily on theEocene and Cretaceous and active collectingof Eocene rocks has provided important in-formation regarding the timing of introduc-tion of several families and genera to NewZealand. With thc exception of LaeviraninaLcirenthey, | 9 29, R h a c' h i o s o mct Woodward,1871, and I-umidocarci nus which werc not-ed by Glaessner ( 1960), all other Eoccne andCretaceous records (Fig. l) have been re-cently described (Glaessner, 1980; Feld-mann and Maxweli. 1990). Thus, Lyreidusde Haan, 1941,l, lotornithrax Griff in, 1963,Leptomithra.r Miers, 1876, and Carcino-plax Mllne Edwards, 1852, extended thcranges of those taxa into the Eocene andHomolodromia l l r lne Edwards, 1880,Hemioon Bell, 1863, Torltnomma Woods,1953, and Eodorippe Glaessner. 1980, werefirs1 records for the Cretaceous. The rangeof Canc'er has been extended with reason-able certainty into the Oligocene and a muchmore questionable occurrence in the Eocenehas also been attrjbuted to the genus. Ca-lappilia sp. has been recently noted fromMiocene rocks (Feldmann and Keyes, 1992),resulting in the first published record ofthatgenus, and the Calappidae, from New Zea-land.
The implications of thesc discoveries aremuch broader than simply extending geo-logical ranges of taxa. Prior to the Oligo-cene. New Zealand occupied a substantiallv
different paleogcographic and paleoccano-graphic setting than it did in post-Eocenetime. Alt"hough details may vary, plate tec-tonic reconstructions of the southern hemi-sphere (Smith et al., l98l; Firstbrook et al.,1979; Scotese, 1979) p laced Austra l ia andNcw Zealand in closer proximity to Ant-arclica in thc Cretaceous and early Tertiarythan in post-Eocene time. The effect of thisplacement was that oceanic circulation pat-terns were markedly different (Shakleton andKennet t , 1973; KenneI I , 1971). Pr ior to theseparation of Australia and New Zealandlrom Antarctica. no circum-Antarctic cur-rent systcm could be established and thcentire region, including the west coast ofSouth America, was influenced by a generalSouth Pacific circulation pattern. At thattime, dispersal of organisms throughout thesouthern high latitudes would have becnreadily possible. Subsequent to breakup ofthesc land masses, however, the circum-Antarctic circulation was rapidly estab-lished and formed a barrier to high latitudedispersal (F ig. l ) .
Employing this paleogeographic framc-work and incorporating the earlier paleo-b iogeog raph i c wo rk o f F lem ing (1962 .1979), Zinsmeister ( 1982) proposcd the ternrWeddell ian Province to reflect the close rc-lationship of pre-Oligocene molluscan as-semblages in New Zealand, Antarctica, andsouthern South America. Following thebrcakup, these areas were isolated from oneanother and followed separate biotic his-tories. T'his work clearly established the im-pact of changing oceanic circulation pat-terns on the molluscan fauna and serves asthc l iamework against which subsequentwork has been cast.
Interpreting the fossil record as i1 was thenunderstood, Fleming (1919) suggested thaltlrc New Zealand decapod fauna was estab-lished in the Miocene or later, somewhatafter most of the molluscan fauna. and con-sequently the affinit ies of the fauna wcrelargely with the Indo-Pacific and Australia.That interpretation, which all ied the NewZealand decapod fauna with Tethyan lau-nas, was consistent with Mioccne pale-oceanographic circulation patterns. How-ever. with the documentation of Eocenc orearlier arrival of decapods in se ven families,including five genera with modern descen-dants. this framework is now untenable. In-
F E I - D M A N N A N I ) N l . L A Y : G E O L O G I C A T - H I S T O R Y O F N E W Z E A L A N D B R A C H Y L J I I A N S 451
O M AH OLOC EN E
l\
B E L L I I D A ED R O M I D A I
G E R Y O N I D A ETlYMENOSOMATIDAILATREILLI DAE
L E U C O S ] I D A EPINNOTHERIDAEC Y M O N O M I D A E
C A N C R ] D A E ?
Fig. I . Paleogeographic maps of the southern hemisphcre showing the positions of continents 80, 40, 20, and
0 million years bcfore present. The oldcst rccords of brachyuran lamilies in New Zcaland are listed on theapproprialc map. Map base from Smith et a/. (198 l).
stead, it appears l ikely that many of the el-ements of the modern New Zealand decapodfauna were introduced in the early Tertiaryfrom high latitude, southern hemisphere ar-eas. Thus, the fauna would contain a strongcomponent of Fleming's Paleoaustral Ele-ment. The geological history of several ofthe families serves to i l lustrate this pattern.
The fossil record of the Homolodromi-idae is restricted to the high southern lati-tudes. As mentioned above, two species intwo genera, including Homolodromia, havebeen described previously from Antarctica(Fcirster et al., 1985: Feldmann and Wilson.
1988) from Miocene and Eocene occur-rences, respectively. The recent recognitionthal. Homolodromia is present in Creta-ceous rocks of New Zealand reinforces thisdistributional pattern and the interpreta-tion, f irst proposed by Feldmann and Wil-son (1988), that this was the area of originof the family. B6ez and Martin (1989) notedthe largely southern hemisphere distribu-tion of l iving species within the genus andarrived at the same conclusion.
In the Raninidae, the paleobiogeographicpattern of Lyreidus has been well docu-men ted (Fe ldmann , 1990 , 1992 ) . The
southern high latitudes are, again, viewedas the region of origin of the genus. Thepresence of Eocene representatives of bothsubgenera within the genus Lyreidus (Lyr-eidus\ bennell i Feldmann and Maxwell,1990, and l,. (Lysirude) v,aitakiensisGlaessner, 1980, i l lustrate the importanceof the New Zealand region in the evolutionof this genus. Newman ( 199 1) noted the dis-tributional pattern of extant and fossil lyr-eidus. particularly emphasizing the presenceof Lyreidus alsearuts Rathbun, 1932, in thenortheast Pacific (Feldmann. 1989) andconcluded that the genus has a Tethyan or-igin. However, the robust Eocene history ofthe genus in the high southern lalitudesstrongly argues for its origins in that regionand subsequent, but early, dispersal of oneofthe l ineages into the northern hemisphere(Feldmann, 1992).
Laeviranina. which is also known fromthe Eocene of New Zealand, may illustratea similar pattern, although the details arenot as well known. This genus, known fromthree species in the New Zealand Eocene,L. perarmata Glaessner, 1960, L. porctrari-ensrs Glaessner, 1980, and L. keyesi Feld-mann and Maxwcll, 1990, has a broad dis-tribution in North America and Europe inthe Tertiary. Feldmann et al. (1993) haverecently described a new species, based uponseveral specimens from Late Cretaceousrocks on James Ross Island, which they re-fer to a new genus ancestral ro Laevirctnina.Thus, the paleogeographic position of theNew Zealand members of this latter genussuggest that they may be parl of anotherexample of high southern latitude origin ofcrabs. This l ineage is l ikely to be ancestral,in 1urn, to the modcrn Rqninoides MilneEdwards, 1837, which has a largely southernhemisphere, Indo-Pacifrc, west African, andCaribbean distribution (Glaessner, 1960).
The record within the Cancridae is lessconclus ive. Nat ions (1975) monographedthe genus Cancer, postulaling a North Pa-cific origin and subsequent radiation. Heconcluded that the sole living species in NewZealand, Cancer novaezelandiae (Jacqui-not, 1853), was rnost closely related to C.edwardsi Bell, 1835, which is known fromthe coast of Peru and Chile. Nations con-cluded that, although there was no fossilrecord of the genus in South America todocument early occurrence on that conli-nent, the New Zealand species was derived
from ancestral South American stock, per-haps in the Miocene. This dispersal patternwould have been in a direction opposite thatofthe prevail ing oceanic current pattern andis considered to be unlikely. Newman ( 199 I )interpreted the distribution of extant andfossil species of Cancer to be an examplc ofreliction of a previously wider ranging Teth-yan complex.
Alternatively. the genus may have bcenestablished in the New Zealand region firstand subsequent ly d ispersed into SouthAmerica. An excellent specimen of Cancernovaezelandiae is known from the Mioceneof New Zealand (Feldmann and Keyes,1992). This occurrence extends the recordof the genus in New Zealand into the Mio-cene. without question. The fossil record ofCancer in pre-Miocene rocks is moreskctchy. An isolated single claw and a badlyeroded carapace, collecled from the base ofthe late Oligocene Cobden Limestone atPoint El izabeth. ncar Greymouth (J3l /f9619), were questionably refcrred to thegcnus (Feldmann and Keyes, 1992). Reex-amination of the material confirms theidentif ication of the claw; howcver, certainaspects oflhe thoracic sternum suggest thatit may not be a (lancer, but, instead, maybe referable to some othcr, as yet undeter-mined, genus. Nonetheless, the claw pro-vides some evidence, albcit sl im, to docu-ment extension of the range of Cancer inlothc Oligocene, the oldest known occurrenceofthe genus. The Eocenc occurrence ofthegenus (Feldmann and Keyes, 1992) is larmore questionable. Thal determination wasbased upon a small chela and two smallf ingers that exhibit some aspects of the mor-phology of Cancer, but differ in others. Un-ti l more complete material is collected, nofirm conclusions should be drawn on thebasis of this material. Therefore. on the ba-sis of the fossil record of Cttnccr, as it is nowknown, the genus may have originated inthe high southern lalitudes and dispersednorthward in a pattern similar to those de-scribed above. The modern restriction ofthe genus to water tempcratures less that25'C (Nations, 1975) is compalible with thisinterprelation.
Analysis of Bias in the Record
As is always the case, there is some dif-ficulty in relating fossil and Recent taxa ow-ing to the lact that different workers nec-
F E L I ) M A N N A N D l l l c L A Y : a i E O L O G I a A L H I S T O R Y O F N E \ \ , Z E A L A N D B R A a H Y t T R A N S
C ARAPAC E WIDTH
453
essarily apply different taxonomic criteriato l iving and fossil material. In addition.these problems are exacerbated because thework usually is done by different individ-uals. Although these problems can never becompletely eliminaled. an attempt has beenmade to minimize the effects by analysis ofrecent summary works and coauthorship ofthe summary results. Unfortunately, thereremains a lack of faunal summaries, for ar-eas comparable to New Zealand. as a basisfor comparison.
2 O c m
2 O c m
-lhe fossil and Recent record ofdecapods
will increase with additional collecting.However, both are moderately well knownand the close correspondence between thetwo suggests that patterns of origin and dis-persal can be discerned. Although the shal-low-water, inshore fauna is extremely wellknown in New Zealand. continued collec-tion in deeper water regions wil l doubtlessprovide additional records at the genus andspecies level, although there is far less like-l ihood that new family records wil l be forth-
Fig. 2- Size lrequency histograms of largest carapace width known from living (upper diagram) and fossil(lower diagram) crabs in New Zealand. The cross-hatched arcas on thc upper diagram ..p..i.nr iaxa knownonly l rom inshore, typical ly rocky. habi tats in waler depths less than l0 m.
454 I O L ] R N A L O F C R T ] S ] ' A C E A N B I O L O G Y . V O L . I 3 . N O . ] . 1 9 9 3
coming. It is far more likely that additionalinformation can be obtained from the fossilrecord. Recent, relatively intensive collect-ing in Eocene and, to a lesser extent, Cre-taceous rocks has yielded significant results.Interest in collecting fossiliferous Mioceneexposures continues, suggesting that this partof the fossil record may be moderately wellknown. However. there has been litt le re-cent collecting in Pliocene and Pleistocenerocks. It is highly likely that, if this part ofthe sequence were investigated in detail, ad-ditional discoveries would be made.
As has been mentioned above and notedby other workers (Bishop, 1986), there issome bias in the fossil record relative to thebody size and ecological preference ofcrabs.In order to assess the extent ofthis problemwith regard to New Zealand fossils, size-frequency histograms of maximum cara-pace width were constructed for living andfossil species (Fig. 2). The sizes plotted weretaken from the published literature, citedelsewhere within the paper, so that each spe-cies is represented within a single size class.A comparison of the two plots indicatesrather close correspondence between livingand fossil size distributions, except in thesmallest size category. The difference in thesmall size fraction is considerably reducedwhen species inhabiting environments lessthan l0 m deep are eliminated. This cate-gory includes all but one of the hymeno-somatids and most of the grapsids, familiesthat are either absent or underrepresentedin the lossil record. Therefore, it appearsthat there is little bias in the fossil recordthat can be attributed directly to small sizeof crab species. Most of the bias can be at-tributed to the potential for preservation,corroborating the results of Bishop (1986).
CoNcrusroNs
The fossil and Recent decapod faunas inNew Zealand show a reasonably close cor-respondence which suggests that there is agood data base, within the existing pub-lished literature, for comparison. The faunais generally cool-temperate in character, withforms derived largely from wide-spread andhigh latitude stock. Nine genera and 39 spe-cies are endemic. Although there is someTethyan influence indicated by the presenceof the Dromiidae, Calappidae, Leucosiidae,and Xanthidae, the role of the high latitudes
as a source of modern forms has been un-derestimated previously. The "typical" NewZealand decapod fauna began to be estab-lished in the Eocene during a time intervalin which New Zealand would have beenmuch more strongly influenced by potentialb iot ic in terchange wi th southern SouthAmerica and Antarctica. This pattern andt iming of launal or ig inat ion is consistentwith the paleoceanographic setting of NewZealand-early and ready dispersal withother Southern Hemisphere sites and sub-sequent paleoceanographic isolation.
AcxNowrsrcEMENTS
Raymond B. Manning, Nat ional Museum ofNaturalHistory, Washington, provided useful commenls on apreliminary draft of the manuscript. The work wassubstantially improved by commcnts from Dr. SylvieSccretan, Mus6um National d'Histoire Naturelle, Par-is. This work was conducled while Feldmann was anErskine Fcllow in thc Department of Zoology. Uni-versity of Canterbury, Christchurch, New Zealand, andthc support of that institution is gratefully acknowl-edged. This is Contribution No. 544, Department ofGeology. Kent State University, Ken1, Ohio 44242,U .S .A .
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F F I I ) \ i \ \ N \ N l ) N t ( 1 , \ \ : ( ; F ( ) L ( ) ( ; l ( . \ l H l s i ( r R \ ' ( ) F N F \ \ l L \ 1 . \ \ l ) l l l < . \ ( H \ l l t \ N S
