Salmlerartige

Salmlerartige
	Sternflecksalmler (Pristella maxillaris)
Systematik
Teilklasse:	Echte Knochenfische (Teleostei)
	Clupeocephala
Kohorte:	Ostarioclupeomorpha
Unterkohorte:	Ostariophysi
	Otophysi
Ordnung:	Salmlerartige
Wissenschaftlicher Name
	Characiformes
	Goodrich, 1909

Die Salmlerartigen (Characiformes), auch Salmler oder Salmlerfische genannt, sind eine Ordnung der Knochenfische. Sie umfasst etwa 1900 Arten, u. a. die Piranhas und Neonsalmler. Einige Arten werden als Speisefische genutzt, viele Arten sind aufgrund ihrer Farbenpracht beliebte Süßwasserzierfische.

Verbreitung

Die Salmlerartigen kommen in tropischen Seen und Flüssen in Südamerika, Zentralamerika und in Afrika südlich der Sahara und im Nil (etwa 210 Arten) vor. Dieses Verbreitungsmuster legt nahe, dass sie sich während der Kreidezeit diversifiziert haben.

Merkmale

Die Ordnung der Salmlerartigen ist vielgestaltig, die meisten Arten sind aber von barbenartiger Gestalt, aalähnlich langgestreckte Formen oder abgeflachte Bodenbewohner fehlen jedoch. Die Salmler Amerikas haben vielfältigere Körperformen hervorgebracht als die afrikanischen Salmler. Wie alle Ostariophysi sind die Salmlerartigen insbesondere durch den Weberschen Apparat, der hier einfach gebaut ist, und eine Reihe knöchriger Strukturen zwischen Schwimmblase und dem inneren Ohr gekennzeichnet. Kennzeichen der meisten Characiformes sind eine kleine Fettflosse zwischen Rückenflosse und Schwanzflosse sowie der Besitz von kräftigen Zähnen. Die Bezahnung im Oberkiefer liegt normalerweise auf dem Prämaxillare. Das Maxillare ist meist nicht oder nur schwach bezahnt. Weiterhin können das Pterygoid und Palatinum bezahnt sein. Die Kieferzähne sind mehrspitzig. Bei den Breitlingssalmlern (Curimatidae) fehlen die Zähne im Alter. Der Oberkiefer ist nicht vollständig vorstülpbar, Ausnahmen sind die Keulensalmler (Hemiodontidae) und die Barbensalmler (Prochilodontidae). Schlundzähne sind normalerweise vorhanden, aber nur bei den Engmaulsalmler (Anostomidae) ähnlich spezialisiert wie bei den Karpfenartigen (Cypriniformes). Barteln fehlen, die Anzahl der Branchiostegalstrahlen liegt bei drei bis fünf.

Der Körper ist normalerweise mit Kammschuppen oder kammschuppenähnlichen Schuppen bedeckt, der Kopf ist unbeschuppt. Schuppen fehlen nur bei der südlichsten Art Gymnocharacinus bergii, die auch keine Fettflosse hat. Die Bauchflossen werden von fünf bis zwölf Flossenstrahlen gestützt, die Schwanzflosse hat normalerweise 19 Hauptflossenstrahlen. Die Afterflosse ist kurz oder gemäßigt lang, mit weniger als 45 Flossenstrahlen. Alle Flossenstrahlen sind Weichstrahlen. Bei den Männchen können die Flossenstrahlen der After- und Bauchflossen kleine Häkchen aufweisen. Die erste Hypuralia ist durch eine Lücke vom Wirbelkörper getrennt. Eine solche Lücke fehlt bei den meisten anderen niederen Teleostei. Die Seitenlinie ist manchmal unvollständig. Salmlerartige sind Physostomen, die zweigeteilte Schwimmblase steht mit dem Darm in Verbindung. Sie dient einigen Raubsalmlern (Erythrinus & Hoplerythrinus) als zusätzliches Amungsorgan.

Viele Salmlerartige sind sehr farbenfroh, viele auch silbrig. Die größte Art ist der 1,40 Meter lange Hydrocynus goliath aus dem Kongo. Einige Arten bleiben auch unter einer Länge von drei Zentimeter, die kleinste wird 13 mm lang.

Lebensweise

Viele Salmler sind Schwarmfische. Sie ernähren sich als Fleisch-, Alles- oder Pflanzenfresser. Zu den Fleischfressern gehören die in lockeren Schwärmen lebenden Piranhas, sowie langgestreckte Raubfische, wie die Spindel- und Raubsalmler, die Amerikanischen und die Afrikanischen Hechtsalmler. Der Wimpelpiranha (Catoprion mento), Gnathodolus bidens und Probolodus heterostomus sind Schuppenfresser. Die Distichodontidae-Gattungen Belonephago, Eugnathichthys, Ichthyoborus & Phago ernähren sich hauptsächlich von den Flossen anderer Fische. Zu den Pflanzenfressern zählen der Schwarze Pacu, die Metynnis- und die Myleus-Arten. Nur wenige Salmler betreiben Brutpflege, der Laich wird meist in Pflanzen abgesetzt. Einige Salmlerartige können akustisch, mit Tönen, die durch die mit Hilfe von Trommelmuskeln in Vibrationen versetzte Schwimmblase erzeugt werden, miteinander kommunizieren. Die chorartigen Laute der Prochilodus können auch außerhalb des Wassers wahrgenommen werden und dienen einheimischen Fischern zum Auffinden der Schwärme.

Äußere Systematik

Die Salmlerartigen gehören zu den Ostariophysi zu denen auch die Karpfenartigen (Cypriniformes) und Welsartigen (Siluriformes) gehören. Sie sind die Schwestergruppe der Siluriphysi (Welsartige & Neuwelt-Messerfische) und bilden mit ihnen das Taxon Characiphysi. Ihre phylogenetischen Beziehungen sind in folgendem Kladogramm dargestellt:

Neuwelt-Messerfische und Welse wie Hypancistrus zebra sind die nächsten Verwandten der Salmlerartigen.

Ostariophysi

Sandfischartige (Gonorynchiformes)

Otophysi

Karpfenartige (Cypriniformes)

Characiphysi

Salmlerartige (Characiformes)

Siluriphysi

	Welsartige (Siluriformes)

	Neuwelt-Messerfische (Gymnotiformes)

Innere Systematik

Es gibt etwa 2000 Arten in ca. 270 Gattungen, 24 Familien und zwei Unterordnungen ^[1]:

Unterordnung Citaharinoidei, Afrika, Zähne zweihöckerig, zweite und dritte Postcleithra (Knochen im Schultergürtel) zusammengewachsen, Neuralbogen des vierten Wirbels autogen, Ctenoidschuppen (Cycloidschuppen bei Citharinus), relativ viele Bauchflossenstrahlen.

Blauer Kongosalmler
(Phenacogrammus interruptus)

Prachtkopfsteher (Anostomus anostomus)

Schlußlicht-Drachenflosser (Gnathocharax steindachneri)

Iguanodectes sp.

- Familie Geradsalmler (Citharinidae)
- Familie Distichodontidae
Unterordnung Characoidei, Afrika, Süd- und Mittelamerika.
- Familie Pracht- und Bodensalmler (Crenuchidae)
  - Unterfamilie Prachtsalmler (Crenuchinae)
  - Unterfamilie Bodensalmler (Characidiinae)
- Familie Afrikanische Salmler (Alestidae)
- Familie Afrikanische Hechtsalmler (Hepsetidae)
- Familie Raubsalmler (Erythrinidae)
- Familie Algensalmler (Parodontidae)
- Familie Wolfssalmler (Cynodontidae)
- Familie Keulensalmler (Hemiodontidae)
- Familie Sägesalmler (Serrasalmidae)
- „Anostomoidea“
  - Familie Engmaulsalmler (Anostomidae)
  - Familie Kopfsteher (Chilodontidae)
  - Familie Breitlingssalmler (Curimatidae)
  - Familie Barbensalmler (Prochilodontidae)
- Familie Amerikanische Hechtsalmler (Ctenoluciidae)
- Familie Schlanksalmler (Lebiasinidae)
- Familie Chalceidae
- Familie Acestrorhynchidae
  - Unterfamilie Acestrorhynchinae
  - Unterfamilie Roestinae
  - Unterfamilie Heterocharacinae
- Familie Iguanodectidae
  - Unterfamilie Iguanodectinae
  - Bryconops clade
- Familie Bryconidae
  - Unterfamilie Bryconinae
  - Unterfamilie Salmininae
- Familie Beilbauchsalmler (Gasteropelecidae)
- Familie Triportheidae
  - Unterfamilie Triportheinae
  - Unterfamilie Agoniatinae
  - Unterfamilie Clupeacharacinae
- Familie Echte Salmler (Characidae)

Die wahrscheinlichen verwandtschaftlichen Beziehungen zeigt folgendes Kladogramm ^[1]:

Characiformes

Characoidei

Echte Salmler (Characidae)

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Chalceidae

	Schlanksalmler (Lebiasinidae)

	Amerikanische Hechtsalmler (Ctenoluciidae)

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Wolfssalmler (Cynodontidae)

Parodontidae

Raubsalmler (Erythrinidae)

	Afrikanische Salmler (Alestidae)

	Afrikanische Hechtsalmler (Hepsetidae)

Pracht- und Bodensalmler (Crenuchidae)

Citaharinoidei

	Geradsalmler (Citharinidae)

	Distichodontidae

Fossilüberlieferung

Das älteste Fossil eines salmlerähnlichen Fisches stammt aus dem Albium, die chronostratigraphisch höchste Stufe in der Unterkreide. Santanichthys aus Brasilien war wahrscheinlich marin oder lebte in Brackwasser und ist wahrscheinlich der älteste Salmlerartige oder der älteste Otophysier. Weiter fossile Salmlerartige sind Sorbincharax aus der ausgestorbenen Familie Sorbincharacidae, Paleohoplias und Tiupampichthys aus Südamerika, Eocitharinus, möglicherweise ein Vertreter der afrikanischen Unterordnung Citaharinoidei und Mahengecharax, möglicherweise die Schwesterart der Alestidae. Die rezente Gattung Brycon ist schon aus dem Oligozän bekannt, Tetragonopterus und Triportheus seit dem Miozän ^[2].

Literatur

Kurt Fiedler: Lehrbuch der Speziellen Zoologie (2. Band, 2. Teil: Fische). Gustav Fischer Verlag., Jena 1991, ISBN 3-334-00339-6.
Joseph S. Nelson: Fishes of the World. John Wiley & Sons, 2006, ISBN 0-471-25031-7.
Günther Sterba: Süsswasserfische der Welt. Urania Verlag., Stuttgart 1990, ISBN 3-332-00109-4.
Petru Banaescu: Zoogeography of Fresh Waters. Aula Verlag., Wiesbaden 1990, ISBN 3-89104-480-1.

Einzelnachweise

↑ ^1,0 ^1,1 Claudio Oliveira et al.: Phylogenetic relationships within the speciose family Characidae (Teleostei: Ostariophysi: Characiformes) based on multilocus analysis and extensive ingroup sampling. BMC Evolutionary Biology 2011, 11:275 doi:10.1186/1471-2148-11-275
↑ K. A. Frickhinger: Fossilien Atlas Fische, Mergus-Verlag, Melle, 1999, ISBN 3-88244-018-X

Weblinks

Commons: Salmlerartige – Sammlung von Bildern, Videos und Audiodateien

Salmlerartige auf Fishbase.org (englisch)
Tree of Life Web Project - Characiformes. Tetras, piranhas, hatchetfishes, headstanders, pencilfishes, and their relatives.
Salmler-Übersicht

[Oliveira_et_al-1] 1,0 ^1,1 Claudio Oliveira et al.: Phylogenetic relationships within the speciose family Characidae (Teleostei: Ostariophysi: Characiformes) based on multilocus analysis and extensive ingroup sampling. BMC Evolutionary Biology 2011, 11:275 doi:10.1186/1471-2148-11-275

[2] K. A. Frickhinger: Fossilien Atlas Fische, Mergus-Verlag, Melle, 1999, ISBN 3-88244-018-X

[1]

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