#NEXUS [*]
[MacClade ToL #28 registered to Proto4 version #28.92   30 March 2001, DO NOT DISTRIBUTE]

BEGIN DATA;
DIMENSIONS  NTAX=2 NCHAR=1;
FORMAT MISSING=? GAP=- ;
MATRIX
[                 ]
[                 ]

Octopoda         ?
Vampyromorpha    ?
;
END;

BEGIN INTERNET;
	NETLINK  TAXON=1  REPLACE LOCATION='?/?.html';
	NETLINK  TAXON=2  REPLACE LOCATION='?/?.html';
END;

BEGIN ASSUMPTIONS;
	OPTIONS  DEFTYPE=unord PolyTcount=MINSTEPS ;
END; 

BEGIN TreeOfLife;
	CladeName Octopodiformes;
	Title 'Octopodiformes_Berthold_and_Engeser,_1987';
	Subtitle Vampire_Squid_and_Octopods;
	Author Name=Richard_E._Young Email=ryoung@hawaii.edu Institution=_ HomePage=_;
	Author Name=Michael_Vecchione Email=vecchione.michael@nmnh.si.edu Institution=_ HomePage=_;
	IndexURL '../../alphabetIndex.html';
	LeadText 'The Octopodiformes contains two orders.';
	LinkedPages 'Index to Cephalopod Taxa=../../cephIndex.html$Index to Cephalopod Families and Genera=../../alphabetIndex.html$Glossary=../../glossary/Glossary.html';
	TitleGraphic '../Vampire.jpg';
	TitleGraphic Ocyanea.jpg;
	TitleGraphicCaption 'Photograph of <B>V. infernalis</B> taken off Southern California (Photograph copyright © 1999, Brad Seibel) and <B>Octopus cyanea</B>,Great Barrier Reef, Australia (photograph copyright © 1999, Mark Norman).


<H4>Other illustrations</H4>

Origin of the Octopoda drawing modified from Young, Vecchione and Donovan, 1999. ';
	IconFolder '/tree/icons/';
	Enclosing Coleoidea;
	EnclosingURL '../coleoidea.html';
	PrevButtonURL '../decapodiformes/decapodiformes.html';
	WebBrowserCreator MOSS;
	Copyright Date=1999;
	ToolBarButton SHOW TOP GIF=indexPhylo.gif ALTGIF=indexPhyloDim.gif URL='../../cephIndex.html' NAME=phylogenetic_index;
	ToolBarButton SHOW TOP GIF=glossary.gif ALTGIF=glossaryDim.gif URL='../../glossary/Glossary.html' NAME=Glossary;
	TEXTNOTE ID=999 TITLE=Introduction TEXT='The Octopodiformes consists of the Octopoda which contains over 200 species and the Vampyromorpha which contains a single species. The latter is a phylogenetic relict.';
	TEXTNOTE ID=1001 TITLE=Discussion_of_Phylogenetic_Relationships TEXT='<TABLE>
<TR>
<TD>Naef (1923) felt that the origin of the Octopoda would "forever be obscure."  But Naef was unaware of the significance of <B>Vampyroteuthis</B> which was believed to be a cirrate octopod at the time. We now know that the <B>Vampyromorpha</B> is the sister group of the Octopoda and that <B>Vampyroteuthis</B> provides telling clues to the origin of the octopods.  Young, <I>et al.</I> (1999) suggest two possible methods for this origin as illustrated below.  In one (benthic route) the pelagic ancestor becomes benthic in a horizontal attitude (similar to the sepiolids today) and subsequently the arms and mouth rotate under the head . The other alternative (pelagic route) the pelagic ancestor develops an "oral" orientation here pictured with the oral end downward (as in <B>Vampyroteuthis</B> today) and subsequently becomes benthic with the oral end down. </TD> 
<TD><IMG SRC="Settlement2.JPG"></TD>
</TR>
</TABLE>

Evidence for the latter route is found in the structure of the brain. Octopods have a series of lobes in the brain, the inferior frontal lobe system, that is derived from the posterior buccal lobe.  These lobes process complex chemotactile information from the arms.  <B>Vampyroteuthis</B> has an "insipient" inferior frontal lobe system (J. Young, 1977).  This bathypelagic animal apparently has a relatively advanced system for processing chemotactile information from the arms, one that surpasses that of shallow-living benthic decapods.  Apparently <B>Vampyroteuthis</B> uses its arms in an unusual manner. Young, <I>et al.</I> (1999) suggest that <B>Vampyroteuthis</B> like its immediate ancestor (a "pre-octopod"), associates with pelagic jellyfish or tunicates and uses its arms and suckers to adhere to and/or explore the surfaces of these gelatinous animals.  That is, it has an oral orientation for contact with surfaces. The pre-octopod, then was pre-adapted for settling on the ocean floor in an oral-end down configuration.  This oral orientation enabled octopods to become the effective crawling animals that most are today. However the manner in which <B>Vampyroteuthis</B> actually uses its arms and its possible association with gelatinous animals has yet to be demonstrated.
<P>
Young, <I>et al.</I> (1999) argue, therefore, that the benthic habitat was the primitive one within the Octopoda and that pelagic species, which comprise nine of the eleven octopod families, are secondarily pelagic. In the Cirrates, these authors point out the compaction of viscera, loss of jet propulsion, presence of a fully-formed inferior frontal lobe system, single oviduct, fusion of the head and mantle and reduction of the shell as evidence of a quasi-benthic ancestry.  In the Incirrates, they point to the presence of corneas (or their remnants), absence of shell, fins and cirri, fully formed inferior frontal lobe system, probably stalked chorions and brooding as evidence of a fully-benthic ancestry.

';
	TEXTNOTE REFINDENT ID=1003 TITLE=References TEXT='Berthold, T. and T. Engeser (1987). Phylogenetic analysis and systematization of the Cephalopoda (Mollusca). Verh. Naturwiss. Ver. Hamburg, 29: 187-220.

Boletzky, S. v.  1992.  Evolutionary aspects of development, life style, and reproductive mode in incirrate octopods (Mollusca, Cephalopoda).  Revue suisse Zool. 99(4):755-770.

Engeser, T. and K. Bandel (1988). Phylogenetic classification of coleoid cephalopods. In: Wiedmann, J. and Kullmann, J. (Eds.), Cephalopods. Present and past. p105-115. Stuttgart

Fioroni, P. 1981. Die Sonderstellung der Sepioliden, ein Vergleich der Ordnungen der rezenten Cephalopoden. Zool. Jb. Syst., 108: 178-228.

Young, R. E., M. Vecchione and D. Donovan. 1999. The evolution of coleoid cephalopods and their present biodiversity and ecology.  South African Jour. Mar. Sci. (in press).';
	TEXTNOTE ID=1 TITLE=Distribution TEXT='';
	TEXTNOTE ID=2 TITLE=Nomenclature TEXT='A variety of names have been proposed for this group  (e.g., Octobrachia, Fiorini, 1981; Octopodiformes, Berthold and Engeser, 1987; Vampyromorphoidea, Engeser and Bandel, 1988; Vampyropoda, Boletzky, 1992). Young, et al.(1999) concluded that Octopodiformes was the most appropriate.';
	TEXTNOTE ID=4 TITLE=Characteristics TEXT='
<OL>
<LI>Arms
<OL TYPE=e>
<LI>Arms II modified or absent.
<LI>Arms IV unmodified.
<LI>Suckers radially symmetrical.
<LI>Horny rings in suckers absent.
<LI>Arms IV united by web (absent in some argonautoids)
</OL>

<LI>Buccal crown - Absent

<LI>Head
<OL TYPE=e>
<LI>Outer statocyst capsule present.
<LI>Superior buccal lobes adjacent (fused at edges) or completely fused to posterior buccal lobes.
</OL>

<LI>Fins
<OL TYPE=e>
<LI>When present, with cartilagenous axial support (only juvenile fin of <B>Vampyroteuthis</B>).
</OL>

<LI>Viscera
<OL TYPE=e> 
<LI>Nidamental glands absent.
<LI>Oviducal glands radially symmetrical.
<LI>Digestive gland duct appendages fused and compact.
<LI>Digestive gland duct appendages lie "outside" nephridial coelom
</OL></OL>
';
	TOPICSORDER 999 4 1001 2 1 1003 ;
	OPTIONS  WRITEASLIST SQUARECORNERS FOOTPARENTHESES FEETINTREE UNDERCONSTRUCTION ExplicitPrevious UPARROWS DEFRANDICON CenterBranches WRITETXPINTT EXCLUDETOLQUERY BREAKAFTERTITLE WRITENEXTPREV NOUSENEXTPREV SPECIFIEDNEXTPREV NUMRANDPICTURES=6 BRANCHHEIGHT=3 INCERTAEEDGEGAP=5 INCERTAETIPLENGTH=6 TAXONSPACING=1;
END;




BEGIN MACCLADE;
Version m.1;
LastModified -1223382951;
Singles 1000&/00;
FileSettings editor ;
Editor 010011 24 Geneva 9 100 1;
TaxStatus closed Geneva 9 U ;
CharStatus closed Geneva 9 U ;
NodeStatus closed Geneva 9 U ;
TreeStatus closed Geneva 9 U ;
TypeStatus closed Geneva 9 U ;
END;