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An Updated Classification of the Recent Crustacea

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DECAPODA: CARIDEA<br />

I <strong>of</strong> course must strongly disagree with <strong>the</strong> proposed<br />

arrangement <strong>of</strong> <strong>the</strong> caridean families into superfamilies,<br />

because I see this as a retrocess from<br />

taxa sustained by apomorphic characters (Christ<strong>of</strong>fersen,<br />

1990) back to groupings based on overall<br />

resemblance, authority (Chace, 1992; Holthuis,<br />

1993), or arbitrary usage. It is true that my proposals<br />

have had little following in <strong>the</strong> carcinological<br />

community, and that some <strong>of</strong> my employed<br />

characters may be questionable. But it is also true<br />

that my efforts remain <strong>the</strong> first attempt to produce<br />

a phylogenetic system <strong>of</strong> <strong>the</strong> Caridea. Because my<br />

system differs substantially from <strong>the</strong> traditional arrangements,<br />

my suggestions have usually been dismissed<br />

as totally heretical, without any serious attempt<br />

to argue alternative possibilities sustained by<br />

better uniquely shared characters. It is ra<strong>the</strong>r depressing<br />

to note that <strong>the</strong> present authors follow this<br />

same tactic. They do not accept a single superfamily<br />

as syn<strong>the</strong>sized in Christ<strong>of</strong>fersen (1990). More<br />

explicitly, but without justification, <strong>the</strong>y reject my<br />

proposal to combine alpheoids, crangonoids and<br />

pandaloids into a monophyletic taxon. This is surprising<br />

to me, because <strong>the</strong>se superfamilies, as redefined<br />

in my cited works, share a remarkable synapomorphy,<br />

<strong>the</strong> multiarticulated carpus <strong>of</strong> <strong>the</strong> second<br />

pereiopod, which is a unique adaptation within<br />

<strong>the</strong> carideans for body cleaning. For this transformation<br />

series, <strong>the</strong>re is even a transitional stage represented<br />

by <strong>the</strong> nematocarcinoids, in which <strong>the</strong> carpus<br />

<strong>of</strong> <strong>the</strong> second pereiopods is longer than in <strong>the</strong><br />

preceding carideans, before being subdivided in <strong>the</strong><br />

sister group represented by pandaloids, crangonoids,<br />

and alpheoids. At a still higher level <strong>of</strong> generality,<br />

this transformation series is congruent with<br />

<strong>the</strong> presence <strong>of</strong> a well developed incisor process on<br />

<strong>the</strong> mandible <strong>of</strong> palaemonoids and all <strong>the</strong> previously<br />

mentioned superfamilies. Going to a lower<br />

hierarchical level, <strong>the</strong>re is fur<strong>the</strong>r congruence with<br />

<strong>the</strong> uniquely expanded first cheliped in crangonoids<br />

and alpheoids. My rearrangements <strong>of</strong> <strong>the</strong> traditional<br />

families into superfamilies eliminate all <strong>the</strong> paraphyletic<br />

family-level taxa, including <strong>the</strong> notably<br />

unsatisfactory Hippolytidae. Finally, just to mention<br />

one remarkable autapomorphy justifying one<br />

<strong>of</strong> my new superfamilies, only palaemonids and<br />

rhynchocinetids share a second distolateral tooth<br />

on <strong>the</strong> basal segment <strong>of</strong> <strong>the</strong> antennule, in addition<br />

to <strong>the</strong> usual stylocerite. Some researchers complain<br />

that I presented few characters for each node, but<br />

this is because my approach is qualitative and I selected<br />

<strong>the</strong> best possible evidence from detailed studies<br />

<strong>of</strong> <strong>the</strong> total morphological and species diversity<br />

<strong>of</strong> <strong>the</strong> Caridea. To refute <strong>the</strong> phylogenetic system,<br />

it is necessary that researchers argue for alternative<br />

replacement characters where <strong>the</strong>y believe I have<br />

failed. Simply ignoring <strong>the</strong> system does not justify<br />

<strong>the</strong> usual assumption that my arrangements are totally<br />

wrong!<br />

Submitted by Martin L. Christ<strong>of</strong>fersen,<br />

Federal University <strong>of</strong> Paraíba, Brazil<br />

DECAPODA: REPTANTIA<br />

I really do not understand why you do not use a<br />

separate category for Reptantia. It is one <strong>of</strong> <strong>the</strong><br />

clearest, most universally accepted groups (taxonomically<br />

or cladistically) among <strong>the</strong> decapods that<br />

we have.<br />

Submitted by Frederick R. Schram,<br />

Zoölogisches Museum, Amsterdam<br />

DECAPODA: ASTACIDEA<br />

You really do get yourselves into deep water when<br />

you try to <strong>of</strong>fer editorial comments on cladistic<br />

analyses. Here you hit ano<strong>the</strong>r one. Where do you<br />

get <strong>the</strong> idea <strong>of</strong> ‘‘extremely primitive Neoglyphea’’<br />

from? Forest and de St. Laurent (1989, Nouvelle<br />

contribution a la connaissance de Neoglyphea inopinata<br />

a propos de la description de la femelle adulte,<br />

Res. Camp. Musorstom 5, Memoirs Mus. Nat.<br />

His. Nat., series A, 144:75–92) made [a] good argument<br />

for allying glypheoids with astacids—not a<br />

particularly primitive alliance. My own preliminary<br />

examination <strong>of</strong> decapod phylogeny (submitted, Hydrobiologia)<br />

not only fairly well confirms <strong>the</strong><br />

Scholtz and Richter scheme, but also squarely places<br />

Neoglyphea within <strong>the</strong> Fractosternalia.<br />

As I say, my own examination <strong>of</strong> <strong>the</strong> subject in<br />

connection with an assignment to address decapod<br />

phylogeny in connection with <strong>the</strong> beginning revision<br />

<strong>of</strong> <strong>the</strong> decapod section <strong>of</strong> <strong>the</strong> Treatise on Invertebrate<br />

Paleontology has, to my surprise, uncovered<br />

<strong>the</strong> basic robustness <strong>of</strong> <strong>the</strong> Scholtz and Richter<br />

analysis. I think you would do well to leave yourself<br />

an opening here.<br />

Of course, I see why you are keen to downplay<br />

Scholtz and Richter because here you adapt a very<br />

conservative combination <strong>of</strong> ‘‘clawed lobsters.’’ I<br />

can accept this for now. However, I think it would<br />

only be fair for you to point out that Scholtz and<br />

Richter would segregate <strong>the</strong> ‘‘clawed (true) lobsters’’<br />

as Homarida from <strong>the</strong> crayfish as Astacida.<br />

My own on-going, recent work indicates that at<br />

least <strong>the</strong> genus Neoglyphea is a fractosternalian in<br />

some kind <strong>of</strong> proximity to <strong>the</strong> Astacida, and that<br />

Enoplometopus may even be a separate clade from<br />

<strong>the</strong> Nephropoidea. That this paraphyly should<br />

emerge among ‘‘lobsters’’ is not too surprising,<br />

since we discover again and again that supposedly<br />

robust, traditional groups bearing a lot [<strong>of</strong>] plesiomorphies<br />

emerge on closer examination as paraphyletic<br />

taxa. Why should macrurous lobsters be<br />

any different?<br />

Submitted by Frederick R. Schram,<br />

Zoölogisches Museum, Amsterdam<br />

DECAPODA: ANOMURA<br />

I fully agree with <strong>the</strong> different parts <strong>of</strong> my specialty<br />

(<strong>An</strong>omura). I agree with <strong>the</strong> changes included in<br />

this new version. As you mention . . . we need more<br />

studies (especially molecular) to improve our<br />

110 Contributions in Science, Number 39 Appendix I: Comments and Opinions

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