Zoology 250 Phylogenetic Trees (2005)


(onychopheran)
The PANARTHROPODA
(= arthropods & nearest relatives)

(tree and traits from
Ruppert & Barnes 1994 p. 1052, and
Brusca & Brusca 2003 p. 501, 875)


(tardigrade)

                    ===========O=== Onychophora (velvet worms; fossil Hallucigenia)
                    |
<<=======P==========|      ====T=== Tardigrada (water bears)
                    ===1===|
                           ====A=== TO ARTHROPODA ====================>

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TRAITS SUPPORTING EACH CLADE (** plesiomorphic- a primitive state, not unique to clade):
P= PANARTHROPODA:
a) elongate body of multiple similar segments**
b) ventro-lateral legs (lobopods) with hooked tips
c) must molt to grow
d) chemical makeup of cuticle
e) ectodermal cilia lost
f) extensive hemocoel ('open' circulatory system), coeloms reduced
g) heart with openings to hemocoel (ostia)
h) panarthropodan sensilla
O= ONYCHOPHORA:
a) external segmentation suppressed
b) unique oral papillae
1:
a) jointed appendages with intrinsic muscles
b) tri-partite cerebral ganglia (proto-, deuto-, & tritocerebrum)
c) loss of circular muscle layer
T= TARDIGRADA:
a) loss of heart
b) loss of nephridia
c) claws of anterior legs modified into stylets & stylet supports
d) tri-radiate, myo-epithelial sucking pharynx
A= ARTHROPODA:
a) lateral compound eyes
b) two tagmata (head, trunk)
c) calcification of cuticle
d) fully segmental sclerites (body-wall cuticular plates)
e) each body segment has 2 sclerites (dorsal tergum, ventral sternum)
f) two-part appendages: basal protopod + distal, multi-segmented part (telopod)
g) complete loss of motile somatic cilia/flagella (some sperm retain flagella)
h) unique hemocyanin respiratory pigment
i) 3-unit brain consisting of a protocerebrum, deuterocerebrum and tritocerebrum (Harzsch 2004 Evol. & Dev. 6:303)



(pycnogonid sea spider)

(horseshoe crab)

(true spider)
Phylum ARTHROPODA

(tree and traits from
Brusca & Brusca 1990 p. 485, 538, 589, 598, 659, 683, 691;
taxa in FULL UPPER CASE indicate the four arthropod subphyla)


(diplopod myriapod)

(chilopod myriapod)

(extinct trilobite)


                                         ==A========= Arachnida (spiders, mites, scorpions, etc.)
                                    ==C==|
   ========= CHELICERATA ===========|    ==X========= Merostomata (horseshoe crabs)
   |                                |
   |                                ===P============= Pycnogonida (sea spiders)
   |
<<=|   ===== TRILOBITOMORPHA ======================== (extinct trilobites)
   |   |
   |   |                                       ====== Symphyla (symphylans)
   |   |                                  ==S==|
   |   |                                  |    ====== Chilopoda (centipedes)
   |   |                ===Myriapoda======|
   =T==|                |                 =========== Diplopoda (millipedes)
       |                |
       |  == UNIRAMIA ==|             ==Entognatha=== Collembola (springtails)
       |  |             |             |
       |  |             ==Hexapoda====|            == Thysanura (silverfish and firebrats)
       |  |               (Insecta)   ==Ectognatha=|
       =M=|                                        == Pterygota (winged insects)
          |
          == TO CRUSTACEA ===========================================================>

Back to Zool 250 tree for animal phyla or Protostomia


TRAITS SUPPORTING EACH CLADE (** plesiomorphic- a primitive state, not unique to clade):
CHELICERATA:
a) two tagmata (cephalothorax=prosoma, without distinct head; abdomen=opisthosoma)**
b) mouth lies before segment 1 embryologically
c) segment #1 limbs= chelicerae (i.e., no antennae; may be pincer-like or fangs)
d) segment #2 limbs= palps or pedipalps
C:
a) cephalothorax has a carapace shield
b) 1st or 2nd abdominal segment modified as genital somite
P= Pycnogonida:
a) pre-oral proboscis
b) ovigers (unusual 3rd pair of limbs)
c) abdomen (=opisthosoma) reduced or absent
d) long, 9-segmented walking legs
e) multiple pairs of gonopores on some/all legs
f) loss of compound eyes
A= Arachnida:
a) abdominal limbs reduced, lost, or modified as spinnerets
b) loss of compound eyes
X= Xiphosura:
a) enlarged cephalothorax with large carapace
b) abdominal limbs modified as book gills
c) long, spiked telson
T:
a) mouth lies before segment 2 embryologically
b) segment #1 limbs= antennae
c) 4 pair post-oral head segments with biramous limbs similar to thoracic limbs
TRILOBITOMORPHA:
a) 3 tagmata (cephalon, thorax, pygidium)
b) body dorso-ventrally flattened
c) limbs of post-oral head segments similar to limbs of thorax (all with two branches, biramous?)**
M= Mandibulata:
a) 2 tagmata (head, trunk of many similar segments)**
b) 5 pair head appendages (2 pair antennae, 1 pair mandibles, 2 pair maxillae)
c) body segment cuticle of 4 sclerites (tergum, 2 lateral pleura, sternum)
d) mouth anterior to segment 3 in adult
UNIRAMIA= Tracheata, = Atelocerata:
a) second antennae lost
b) mandible is 'whole-limb'
c) unique tracheal system
d) uniramous limbs (exopodite lost)
Myriapoda:$
a) 2 tagmata (head, trunk of many similar segments)**
b) loss of compound eyes
c) loss of palps on first & second maxillae
d) repugnatorial glands
S:
a) medial fusion of both pairs of maxillae
b) maxillipeds modified as raptorial poison fangs
Hexapoda:
a) second maxillae fuse as labium
b) 3-segment thorax with limbs
c) 11-segment abdomen, no limbs
Entognatha:
a) mouthparts sunk in a pouch
Ectognatha:
a) mouthparts not sunk in pouch
Pterygota:
a) possess wings on 1 or 2 thoracic segments

$ increasing evidence suggests Myriapoda branched off before the Hexapoda and Crustacea diverged (e.g., these two groups both have a 4-part crystalline cone in their ommatidia but myriapods do not; Harzsch 2004 Evol. & Dev. 6:303), so Myriapoda may be a separate subphylum (i.e., subphylum Uniramia is rejected).


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Copyright (c) 2005 by A. Richard Palmer. All rights reserved.
(revised Jan. 5, 2005)