The Vertebrates Novumbra & Esocidae

Euteleostei: Neognathi 

Novumbra & Esocidae

Abbreviated Dendrogram
Teleostei │ └─Euteleostei ├─Ostariophysi │ ├─Anotophysi │ └─Otophysi │ ├─Cypriniformes │ └─Siluriphysi └─Neognathi ├─┬─Salmoniformes │ └─Esociformes │ ├─Umbra │ └─┬─Dallia │ └─┬─Novumbra │ └─Esocidae │ ├─Estesesox │ └─Esox └─┬─Argentiniformes └─Neoteleostei ├─Stenopterygii └─Eurypterygii ├─Cyclosquamata └─Ctenosquamata ├─Scopelomorpha └─Acanthomorpha

   Umbra & Dallia
   Novumbra & Esocidae

Taxa on This Page

  1. Argentiniformes
  2. Esocidae
  3. Esox
  4. Estesesox X
  5. Novumbra


NovumbraNovumbra:  N. hubbsi Schultz 1929. (Olympic mudminnow).

Range: from the Oligocene [K+86] of North America (Washington State, USA)

Phylogeny: Esociformes ::: Esocidae + *.

Characters: up to 8 cm [PB91]; premaxilla borders only 1/3rd of gape [WV82]; premaxilla with prominent articular process on dorsal margin fitting against head of maxilla [WV82]; maxilla elongate, with single supramaxilla [WV82]; frontal comes to a square end over sphenotics [WV82]; frontal with distinct ridge across widest portion [WV82]; deep posttemporal fossa behind ridge [WV82]; frontal posterolateral margin concave [WV82]; frontal overlies anterior 1/3rd of supraoccipital, with sharp angle between posterior & posterolateral margins [WV82]; parietals very small [WV82]; posttemporals large & plate-like (compared to other primitive esociforms) [WV82]; preopercular arms about same size [WV82]; opercular subtriangular, with "wavy" posterior border [WV82]; subopercular large, with prominent anterior process [WV82]; interopercular long, with vertical posterior margin [WV82]; exoccipitals antero-posteriorly short [WV82]; sphenotic without lateral process at apex [WV82]; alisphenoids do not meet on the midline [WV82]; proethmoid entirely dermal, with anterior end expanded & curved laterally [WV82]; Novumbra hyoid arch from [WV82] proethmoid anterior margin not thickened & fits into groove on premaxilla [WV82]; vomer club-shaped, with tapered posterior end ventral to parasphenoid [WV82]; palatine short & stout [WV82]; palatine without anterior facet & with posterior end deep & truncated [WV82]; ectopterygoid large, with anterior & posteroventral limbs [WV82]; ectopterygoid, anterior process fits into posterior of palatine [WV82]; metapterygoid small & without struts [WV82]; symplectic nearly straight [W84]; parasphenoid narrow, waisted & elongate extending from mid-vomer level to mid-basioccipital [WV82]; parasphenoid with slight, ascending alar processes posterior to waist & reaching up to prootics, with second pair of alae sometimes present [WV82]; carotid foramen between parasphenoid & prootic just posterior to alae [WV82]; anterior of parasphenoid concave ventrally [WV82]; lower jaw short & robust, with greatest depth at posterior of dentary [WV82]; dentary, ventral margin excavated anteriorly, with tube for sensory line [WV82]; anguloarticular posterodorsal border inclined anteriorly [WV82]; coronomeckelian very small, nested at mid-length of anguloarticular [WV82]; premaxilla with single row of 5-7 teeth [WV82]; vomer bearing 17-21 recurved teeth (mostly) anteriorly [WV82]; palatine with two rows of teeth with 12 (lateral) & 5 (medial) teeth [WV82]; dentary with single row of 18 recurved teeth [WV82]; hyomandibular shaft long & opercular process short [WV82]; epihyal somewhat rectangular [WV82]; ceratohyal asymmetrical & without processes [WV82]; dorsal hypohyal much smaller than ventral [WV82]; urohyal with weak vertical keel and with anterior sockets for sternohyoideus ligament (from ventral hypohyal -- acts in opening & expanding mouth) [WV82]; branchiostegal rays acinaciform & numbering 7-9, with 2 attached to epihyal [WV82]; 4th epibranchial with uncinate process [WV82]; but 4th external levator (brachialis?) muscle absent [JP97]; branchial toothplates on basihyal & basibranchials 3-4 [WV82]; ceratobranchial 5 short [WV82]; 5th epibranchial absent [JP97]; gill rakers small, finely toothed irregular plates [WV82]; 37-38 vertebrae, with 18-19 precaudal [WV82] ([N94]: 37-40 total); centra mostly longer than wide [WV82]; neural & hemal spines [sic -- probably means "arches"] fused to centra [WV82]; anterior 3-5 neural spines expanded distally, followed by 12-14 spines bearing tubular supraneurals just anterior & parallel to body of neural spine [WV82]; 12-14 all precaudals with pleural ribs attached to separate parapophyseal ossifications [WV82]; dorsal fin above or slightly in front of anal fin origin [PB91]; anal & dorsal fin bases about same length [PB91]; dorsal fin with 12-15 rays [PB91] [N94]; anal fin with 10-13 rays [PB91] ([N94]: 11-13); epineurals present throughout & epihemals (?!?) in caudal region [WV82]; first preural with neural arch plus either 1 epural or a neural spine [WV82] (contra [R74]: neural arch present with single epural); 5-7 hypurals with 1st large and 2-3 slender [WV82]; urodermals don't reach anteriorly beyond first ural centrum [R74] [WV82]; caudal fin straight edged or slightly indented [PB91]; 5-7 procurrent caudals above & below fin , with 18-19 principal caudal rays [WV82]; extrascapular present [JP97]; small coracoid & 1 postcleithrum present [WV82]; 4 proximal pectoral radials & 15-21 fin rays [WV82] (compare [PB91]: 18-25 rays & [N94]: 18-23); pelvic girdle with elongate anterior pubic process & rounded illiac posterior plates, 5-7 fin rays [WV82] (compare [PB91] & [N94]: 6-7 rays); lateral line absent [PB91].

Image: N. hubbsi by Tom Baugh from The Olympic Mudminnow

Links: The Olympic Mudminnow; FISHBASE; Frasier Glaciation.

References: Johnson & Patterson (1997) [JP97]; Kettler et al. (1986) [K+86]; Page & Burr (1991) [PB91]; Rosen (1974) [R74]; Wilson (1984) [W84]; Wilson & Veilleux (1982) [WV82]. ATW030412.

Esox americanus (redfin)Esocidae:Oldmanesox

Range: from the Late Cretaceous (Campanian) of North America, Asia & Europe

Phylogeny: Esociformes ::: Novumbra + * : Estesesox + Esox

Characters: primitively as small as Umbra? [W+92]; long and usually duckbill-shaped jaws [W84]; basisphenoid present [JP97]; palatines broad & oriented anteroposteriorly [W+92]; depressible teeth [W84]; teeth with truncated, C-shaped tooth bases [W+92]; palatine teeth in multiple, parallel, anteroposterior tooth rows extending far posteriorly on wide shelf [W+92]; palatine teeth with C-shaped bases concave posteromedially [W+92]; palatines with anterolateral maxillary & anteromedial ethmoid facets opposite each other & joined by horizontal cleft [W+92]; mandibular sensory canal relatively small (compared to basal esociforms) [W+92]; dentary with low coronoid process [W+92]; dentaries long & slender, bearing median shelf & several rows of depressible teeth anteriorly with C-shaped bases concave posteromedially [W+92]; 4th external levator brachialis?) muscle absent [JP97]; 4th epibranchial with uncinate process, but without levator process [JP97]; body elongate [W84]; fins are soft rayed, lacking spines; dorsal and anal fins located opposite each other about three-quarters of the way toward tail; adipose fin absent; pelvic fins located at mid-length.

Note: see detail of Oldmanesox at left. In point of fact, many of my socks do look like this.

Image: Esox americanus (right) from the Virtual Aquarium. Note the odd resemblance, especially in the skull, to cephalaspids such as Ateleaspis.

Links: The Virtual Aquarium of Virginia Tech--Pikes; Gli Esocidi, Esocidae Italian); Pike Family Esocidae; Artstre- Esocidae Norwegian); Hechte Esocidae) (German); Famille Esocidae (French); The Journal of Vertebrate Paleontology; NAS - Query Results Huge pictures and lots of wildlife management data).

Links: Johnson & Patterson (1997) [JP97]; Wilson (1984) [W84]; Wilson et al. (1992) [W+92]. APW 040806.

Estesesox:E. foxi Wilson et al. 1992.

Range: Late Cretaceous (Campanian to Maastrichtian) of North America.

Phylogeny: Esocidae : Esox + *.

Characters: palatine with prominent longitudinal ridge dorsally [W+92]; palatine anterior margin, anteromedial corner extends further anteriorly [W+92]; palatine with "roofed" facet for maxilla on anterolateral corner & "roofed" facet for ethmoid on medial edge directly medial from maxillary facet [W+92]; dentary with lateral trigeminal foramen located near tooth bases and oriented transversely (not posteriorly -- that is the medial and lateral foramina are located almost directly across from each other) [W+92]; dentary with trigeminal foramina located posteriorly, about at mid-length of dentary, with lateral & medial foramina directly opposite each other on the dentary ramus [W+92]; dentary trigeminal foramina just behind posterior terminus of area with multiple tooth rows [W+92]; dentary ventral margin concave but not excavated [W+92]; no fixed teeth -- all depressible [W+92]; palatines with numerous straight, anteroposterior rows of depressible teeth [W+92]; palatines with C-shaped tooth bases concave posteromedially [W+92]; palatine teeth larger anteriorly & medially except at extreme anterior end [W+92]; dentary with tooth-bearing margin nearly straight to slightly concave (dorsally? laterally?) in lateral view [W+92]; dentary without fixed teeth (all teeth depressible) [W+92]; dentary symphysis deep, with prominent medial tooth-bearing shelf [W+92]; 5- tooth rows near dentary symphysis [W+92]; dentary most medial tooth row longest & continuing to posterior end [W+92].

Note: some of these characters are a bit questionable, given the very scrappy materials. However, the position and orientation of the trigeminal foramina seems to be quite distinctive & diagnostic.

References: Wilson et al. (1992) [W+92]. APW 040806.

Esox lateral skull, suspensorium & dorsal skull [J75].Esox:Linnaeus, 1758.  E. lucius Linnaeus 1758 (pike or northern pike); E. americanus Gmelin 1798 (redfin, grass pickerel); †E. borealis; †E. dispar; †E. kronneri Grande 1999; †E. lepidotus Agassiz 18??; E. masquinongy Mitchill 1824 muskellunge); E. niger Lesueur 1818 (chain pickerel); †E. papyraceus; E. reichertii Dybowski 1869 (Amur pike);  †E. tiemani Wilson 1980 [W80]. Coextensive with living Esocidae.

Range: from the Paleocene, presently found in fresh waters of  Eurasia and North America, usually colder waters.

Phylogeny: Esocidae : Estesesox + *.

Characters:  maximum size 33-165 cm & 200-3200 g; long, flat, duckbill-like snout [C96] [PB91] [N94]; premaxilla small & fixed with connective tissue [J75]; maxilla head with palatine process [WV82] [J75]; single supramaxilla [J75]; supramaxilla extends posteriorly a bit further than maxilla [W84]; nasals present [C96] [N94] (contra [W84]: not found in E. tiemani); nasals tubular [J75]; frontals elongate & narrow anteriorly [W84]; parietals relatively large, meet at midline and extend to pterotics, roofing posttemporal fossa [WV82] (contra [J75]: parietals small & separated at midline by supraoccipital); pterotics elongate [W84]; intercalar present joining pterotic, exoccipital & epidotic at their junction with ventral ramus of posttemporal [WV82]; sphenotic with lateral process at apex for origin of m. levator arcus palatini [WV82]; dermosphenotic present in larger individuals [J75] [3]; orbits moderately large [C96]; orbits with 2 unossified scleral cartilages [J75]; large myodome [JP97]; hyomandibular with long, posteroventral opercular process, lateral preopercular strut, anteroventral flange against metapterygoid, and additional anteroventral process at right angle to opercular arm [W84]; mesopterygoid small [W84]; metapterygoid large [W84]; quadrate robust, with prominent anterodorsal strut supporting ectopterygoid [W84]; symplectic nearly straight [W84]; preopercular with slight bend at the angle, sensory canal along entire length and with anteromedial flange [W84]; subopercular slightly curved, with almost parallel dorsal & ventral margins & prominent dorsal process at anterodorsal corner [W84]; opercular relatively rectangular, but proportions vary [W84]; basioccipital includes first vertebral centrum [J75]; parasphenoid, anterior end dorsal to part of vomer [J75]; parasphenoid expanded under otic capsule, with ascending process associated with spiracle [J75]; parasphenoid narrow & elongate [W84]; alisphenoids meet on the midline [WV82]; proethmoid thickened & grooved posteriorly to cap anterior end of rostral cartilage & articulate via joint with anteromedial margin of palatines (joint may include endochondral ossification) [WV82]; vomers truncated anteriorly [W84]; palatine fused to autopalatine (anterior palatoquadrate derivative) [J75]; ectopterygoid robust, with prominent ethmoid process [W84]; gape large [C96]; dentary a fused mental ossification and probably a fused splenial [J75]; coronomeckelian is small ossification around adductor fossa [J75]; anguloarticular invades Meckelian to form articular surface [J75]; retroarticular from distal Meckelian ossification [J75] [4]; retroarticular includes a dermal plate as well as perichondral ossification of? around? the Meckelian [J75]; branchiostegal rays 9-20 [C96] [W84] ([N94]: 10-20]); branchiostegal rays variably divided between halves of ceratohyal [W84]; teeth on jaws, vomer, palatine & tongue (?!) [C96]; palatine is main tooth-bearing surface of upper jaw [J75]; palatine elongate, with numerous depressible teeth, largest anteromedially [W84]; depressible teeth leaving characteristic truncated, C-shaped root [W84]; dentary strongly toothed, with largest teeth at mid-length [J75]; fixed, canine teeth posterior, depressible teeth anteriorly [W84]; ceratohyal elongate & waisted [W84]; 2 hypohyals, with ventral larger [J75] (ventral not larger in E. tiemani? [W84]); gill rakers reduced to patches of small, very sharp denticles [C96]; lines of small toothplates on hypobranchials 1-3 & ceratobranchials 1-4 [J75]; basihyal & basibranchials 1-2 are ossified, with basihyal joined to basibranchial 1 by toothplate [J75]; hypobranchial 3 extends well below level of basibranchials [J75]; 5th epibranchial absent [JP97]; entoglossal & urohyal present [J75]; body long & cylindrical in younger individuals, laterally compressed in older [C96]; possibly unique "nape bone" as late-developing sliver lying above neural spine of 1st vertebra [J75] [5]; 42-67 vertebrae [C96] [W84] [N94]; caudal vertebrae 18-23 [W84]; neural arches not fused to centra? [JP97]; only first few epineurals fused to arches [JP97]; dorsal & anal fins located far posteriorly [W84] [PB91]; dorsal fin rays 15-26 [W84]; anal fin abdominal [PB91]; anal fin rays 14-22 [W84]; forked caudal fin [C96] [PB91] [N94]; preurals 1-2 may have single neural arch [R74]; 2-3 epurals, with no full neural spine on preural 1 [R74] [W80]; 5-7 hypurals with 1st large and 2-3 slender [WV82] [R74] [W84]; urodermals reach anteriorly beyond first ural centrum [WV82]; 5-7 procurrent caudals above & below fin , with 18-19 principal caudal rays [WV82] (contra? [C96] [N94]: 40-50 rays), with 16-17 branched [R74] [C96] [N94]; no fin spines [PB91]; paired fins small to moderate size [C96]; extrascapular variably present [J75] [W84]; pectoral fin rays 11-18 [W84]; pelvic girdle with elongate anterior pubic process, thin medial pubic plates & rounded posterior iliac processes [W84]; pelvic fin rays 8-13 [W84]; many small cycloid scales [C96] [PB91]; lateral line complete & straight, with notched (heart-shaped) scales on the lateral lines [C96] (contra [W84] no cardioid scales in E. tiemani); air breathing using adaptations of swim bladder [C74]; voracious predators, feeding on a wide variety of fishes and other vertebrates; some tolerance for brackish water; normally a shallow water visual predator [C96].

Note: [1] This genus has a good deal of internal complexity, with several well-known species, races and subspecies, as well as a subgenus. [2] E. tiemani is quite interesting from the point of view of polarity. Some key meristic characters fall toward extremes of the Esox range: 40 + 19 vertebrae (low), 3 epurals (high), 6 hypurals (average), 11-12 branchiostegals (low), length 34 cm. These tend to fall towards the Umbra side of the distribution, although they are still within the range of living Esox except for the third epural) [W80]. [3] [WV82] figure a dermosphenotic in Novumbra but not in Umbra or Dallia. On the other hand, they do not discuss it, and it would be hard to draw any firm conclusion. [4] In other words, the anguloarticular is homologous with only the angular. The retroarticular is homologous only with the articular. If confirmed in other esocids, this would be a rather interesting trait. However Jollie suggests that this condition is not that unusual in teleosts. [5] On the other hand, this could simply be a detached "accessory neural arch" mentioned by [JP97], which is relatively common. The accessory neural arch may, in turn, be a displaced neural arch belonging to the centrum fused to the basioccipital. Alternatively, it may be a supraneural associated with the same vertebra -- or half a dozen other possibilities. This whole anatomical region is so full of vague and uncertain findings (or perhaps just findings we don't understand?) that we've deliberately left a certain amount out in this summary.

Image: note that, in some cases, the images from Jollie (1975) use rather different nomenclature from that used elsewhere in this section. Our apologies for the somewhat crude image, but it saved considerable time & bandwidth.  Compare the less detailed, but more legible, images of more basal esociform skulls in this section for orientation.

Links: Northern Pike; Genera Reference- Detail; Muskellunge Esox masquinongy; Animal Diversity Web: Esox lucius (Northern Pike)- Narrative; The Pike (Best on the Web); AES 44 Cavender abstract).

References: Crawford 1974) [C74]; Crossman 1996) [C96]; Johnson & Patterson (1997) [JP97]; Jollie (1975) [J75]; Nelson (1994) [N94]; Page & Burr (1991) [PB91]; Rosen (1974) [R74]; Wilson (1980) [W80]; Wilson 1984) [W84]; Wilson & Veilleux (1982) [WV82]. APW 040806.

herringArgentiniformes:  Argentina, Dolichopteryx, Bathylagus, Nansenia (herring, smelt, etc.). Argentinoids & alepocephaloids.

Range: from the Late Cretaceous?

Phylogeny: Neognathi :: Neoteleostei + *.

Characters: basibranchial toothplate edentulous and with sharp median ridge of bone & a membrane which divides orobranchial chamber into right & left halves [R74]; "crumenal organ" on 5th ceratobranchial involves an extra cartilage & interdigitating gill rakers -- for filter feeding or grinding) [R74] [$JP97]; unfused 5th epibranchial [R74]; caudal median cartilages present, jointly supporting lowermost ray of upper fin lobe [$JP97].

References: Johnson & Patterson (1997) [JP97]; Rosen (1974) [R74]. APW 040806.

checked ATW060106