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The Place of the Hoplocarida in the Malacostracan Pantheon

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The Place of the Hoplocarida in the Malacostracan Pantheon The University of Maine DigitalCommons@UMaine Marine Sciences Faculty Scholarship School of Marine Sciences 6-1-2009 The lP ace of the Hoplocarida in the Malacostracan Pantheon Les Watling University of Maine - Main, [email protected] C. H.J. Hof F. R. Schram Follow this and additional works at: https://digitalcommons.library.umaine.edu/sms_facpub Repository Citation Watling, Les; Hof, C. H.J.; and Schram, F. R., "The lP ace of the Hoplocarida in the Malacostracan Pantheon" (2009). Marine Sciences Faculty Scholarship. 134. https://digitalcommons.library.umaine.edu/sms_facpub/134 This Article is brought to you for free and open access by DigitalCommons@UMaine. It has been accepted for inclusion in Marine Sciences Faculty Scholarship by an authorized administrator of DigitalCommons@UMaine. For more information, please contact [email protected]. JOURNALOF CRUSTACEANBIOLOGY, 20, SPECIALNUMBER 2: 1-11, 2000 THE PLACE OF THE HOPLOCARIDA IN THE MALACOSTRACAN PANTHEON Les Watling, Cees H. J. Hof, and Frederick R. Schram (LW,corresponding) Darling MarineCenter, University of Maine, Walpole, Maine 04573, U.S.A. (e-mail: [email protected]);(CHJH) Department of EarthSciences, University of Bristol, Wills MemorialBuilding, Queens Road, Bristol BS8 1RJ, United Kingdom (e-mail: [email protected]); (FRS) Zoological Museum, University of Amsterdam,Post Box 94766, NL-1090 GT Amsterdam, The Netherlands(e-mail: [email protected]) ABSTRACT The stomatopodbody plan is highly specializedfor predation,yet the SuperorderHoplocarida originatedfrom something other than the "lean,mean, killing machine" seen today.The fossil record of the groupindicates that it originatedearly on froma non-raptorialancestor, with the specialized predatorymorphology developing much later. The RecentHoplocarida have been variouslyposi- tionedwithin the Malacostraca,from a Subclassequal in rankto the Eumalacostraca(= Caridoida) to being placedas a Superorderwithin the Eumalacostraca.Consideration of the early fossil mor- phology,especially of the form of the carapace,of the positionand functioningof the articlesin the last threepairs of thoracopods,and of otherfeatures, suggests that hoplocarids are earlyderiv- ativesof a basaleumalacostracan stock that was "shrimp-like"in form.The enhancementof an ab- dominalrespiratory system most likely allowed the developmentof the anteriorthorax into the specializedraptorial system present today. Wewant to build a phylogenetictree thatpresents the actual evolutionaryhis- tory of the organisms in it. For phylogeny to be correctly inferredfrom the wealthof data available,it is necessaryto identifyinformative homologous fea- tures that allow us to unite organisms in the tree. Sorting out ho- mology from homoplasy is one of the chief pastimes of phylogeneticists. -Raff (1996) The Hoplocaridaare best known today as Ruppertand Barnes (1994) rank the Hoplo- the Order Stomatopoda, the "lean, mean, carida as a Subclass of the Class Malacos- killing machines" of the shallow marine traca,equivalent in rankto the Phyllocarida world. Yet, while it is obvious that the mor- and Eumalacostraca;Meglitsch and Schram phology embodied in the modem stomato- (1991) move the Phyllocaridato a new Class pods is highly specialized, the origins and Phyllopodaand rank the Hoplocaridaand Eu- affinitiesof that morphologyare not quite as malacostracaas subclasses of the Malacos- clear (e.g., Schram, 1969, 1974, 1986; traca;and Brusca and Brusca (1990) place the Hessler, 1982; Dahl, 1983a; Wheeler, 1998; Hoplocaridaas a Superorderwithin the Eu- Wills, 1998). In this paper, the base of the malacostraca,leaving Phyllocaridaas a sub- hoplocaridanclade will be examinedwith a class of the Malacostraca,essentially as Cal- view to establishing its affinities and taxo- man had done. Among treatmentsdealing nomic relationshipswith other malacostra- strictlywith crustaceans,Bowman and Abele cans. (1982), Schram (1986), and Forest (1994) Hoplocaridahave historicallybeen difficult move the Hoplocaridaout of the Eumalacos- to place. Calman(1909) arrangedthe hoplo- traca, whereas Kaestner(1970) divides the caridansalong with other groups within the Malacostracainto six superorders,one of Series Eumalacostracaalongside the Series which is the Hoplocarida.In the presentpa- Phyllocarida.There seems to be no univer- per, we treat the hoplocarids as the sister sally accepted position for the group today, taxon of the syncarid/eucaridline (Table 1). however. Modem invertebratezoology text- From a paleontologicalas well as neonto- books have taken a variety of approaches: logical perspective, the Hoplocaridais rep- 1 2 JOURNALOF CRUSTACEANBIOLOGY, VOL. 20, SPECIALNO. 2, 20u0 Table 1. Conceptual arrangement of the malacostracan orders with stenopodous thoracic limbs (= Eumalacostraca, sensu lato), after Watling (1983, 1999) and Schram (1986), with modifications according to the suggestions in this paper. Known geological ranges are given. Taxa marked with a '?' are of uncertain rank. This schema is not to be seen so much as a functional taxonomy but as a heuristic device to focus discussions and stimulate more detailed and comprehensive cladistic analyses in the future. There is much to be gained by setting aside facies concepts and focusing on identifying and rigorously defining monophyletic groups. Hoplocarida Palaeostomatopodat (L. Carboniferous-U. Carboniferous) Stomatopoda Archaeostomatopodeat (M. Mississippian-U. Pennsylvanian) Unipeltata (L. Jurassic-Recent) ?Syncarida Palaeocaridaceat (L. Carboniferous-L. Permian) Anaspidacea (Triassic-Recent) Bathynellacea (Recent) "Carida" Decapoda (L. Devonian-Recent) Aeschronectidat (M. Pennsylvanian) Belotelsonideat (Carboniferous) Waterstonellideat (Carboniferous) Euphausiacea (Recent) Pygocephalomorphat (Carboniferous-Permian) Lophogastrida (Pennsylvanian-Recent) Mysida (?Jurassic-Recent) Peracarida Mictacea (Recent) Spaleaogriphacea (Mississippian-Recent) Thermosbaenacea (Recent) Tanaidacea (L. Carboniferous-Recent) Cumacea (?Pennsylvanian-Recent) ?Amphipoda (L. Eocene-Recent) ?Isopoda (M. Pennsylvanian-Recent) resentedby one modem order,the Stomato- Calman(1909) gave a brief history of the poda and the extinct Orders Palaeosto- Malacostracaand then provided definitions of matopoda(Late Devonianthrough Late Mis- the groupshe had proposedfive years earlier sissippian)and Aeschronectida(see Schram, (Calman, 1904). These include the Series 1986). The OrderStomatopoda is subdivided Leptostracaand Eumalacostraca,the latter into two Suborders,the Unipeltata (Upper containing the Divisions Syncarida, Per- Jurassicto Recent) containingfossil taxa as acarida,Eucarida, and Hoplocarida. Calman's well as the modem stomatopods,and the Ar- definitionof the Eumalacostracais as follows: chaeostomatopodea(Middle Pennsylvanian to "Abdomenof six somites (the numbermay UpperPennsylvanian) containing only a small be reducedby coalescence),the last of which groupof extinctforms. Recently, Jenner et al. typically bears a pair of appendages,and a (1998) uncovered the possibility that the telson which never bears moveable furcal Palaeostomatopodamight be paraphyleticrel- rami; no adductormuscle of the carapace; ative to the monophyleticStomatopoda. thoracic limbs rarely all similar (Euphausi- acea), typicallypediform, protopodite of two ARE HOPLOCARIDSEUMALACOSTRACANS? segments except in Stomatopoda"(Calman, As a startingpoint, the early definitionsof 1909: 148). Fromthe beginning,then, the def- the Eumalacostracaand the Hoplocaridaneed initionof the Eumalacostracahas exceptions, to be examined.Schram (1969) noted, after butseems to be centeredaround Calman's con- a review of the variouslysuggested phyloge- cept of a "caridoid"facies (Hessler,1982a). netic positions, that the "definitionof a hop- Phyllocarids differ from the other mala- locarid varied from author to author, de- costracans in having polyramous phyl- pendingon the phylogeneticinterpretation he lopodouslimbs andseven abdominalsomites, adopted"(p. 277). one of which is without appendages.Exact WATLINGET AL.: PHYLOGENETICPOSITION OF THE HOPLOCARIDA 3 definitions for the non-phyllocarid malacos- If we focus on hoplocarids and eumala- tracans have always been troublesome be- costracans, Malacostraca sensu stricto are cause of the great diversity of body forms en- united by the following features: "naupliar compassed by all the orders, both living and eye of three cups each with three everse sen- fossil. One solution, derived from the cladis- sory cells, polyramous and stenopodous tho- tic analysis of Schram (1986), was to move racopods, uropods, and a postcephalic cara- the phyllocarids out of the Malacostraca and pace structure that does not envelop the ab- unite them with other phyllopodous limb- domen or thoracic limbs" (Schram, 1986: bearing groups such as the cephalocarids and 528). From there Schram noted that eumala- branchiopodans. This plan generated another costracans have a uniarticulate antennal scale suite of criticisms, but it had the advantage of and strongly elaborated abdominal muscula- making the Malacostraca more homogeneous. ture to facilitate the caridoid escape reaction, By 1969, the effective definition of Eu- while hoplocarids possess triramous anten- malacostraca had become generalized to in- nules, three-articulate thoracopod protopods, clude all malacostracans except phyllocarids: four-articulate thoracopod outer branch, at "Malacostraca generally of shrimp-like form least in the fossil forms, (if one accepts the distinguished from the Phyllocarida
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