Figure 1.
Juvenile Stage I of Austropotamobius torrentium. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D-H) pereiopods 1-5, dorsal view or lateral view; (I-M) distal segments of pereiopods 1-5, dorsal view or lateral view. Scale bars: 0.25 mm.
Figure 2.
Juvenile Stage I of Austropotamobius torrentium. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. bsg, basal segment of peduncle; bsp, basipodite; c, carpopodite; cxp, coxopodite; d, dactylopodite; de, distal endite; dsg, distal segment of peduncle; ed, endopodite; en, endite; ep, epipodite; exp, exopodite; exf, external flagellum; i, ischiopodite; in, incisor ridge; inf, internal flagellum; m, meropodite; mp, mandibular palp; mr, molar ridge; msg, intermediate segment of peduncle; p, propodite; pe, proximal endite; tc, toothed crest. Scale bars: 0.25 mm.
Figure 3.
Juvenile Stage II of Austropotamobius torrentium. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D-H) pereiopods 1-5, dorsal or lateral view; (I-M) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 4.
Juvenile Stage II of Austropotamobius torrentium. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) flagellum of antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view, (J) pleopod. cd, caudal process; cp, cephalic process; in, incisor ridge. Scale bars: 0.25 mm.
Figure 5.
Juvenile Stage III of Austropotamobius torrentium. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D-H), pereiopods 1-5, dorsal or lateral view; (I-M) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 6.
Juvenile Stage III of Austropotamobius torrentium. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) flagellum of antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm, C: 1.0 mm.
Figure 7.
Juvenile Stage I of Procambarus virginalis. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan and telson thread, dorsal view; (D) posterior edge of the tail fan and telson thread; (E-I), pereiopods 1-5, dorsal or lateral view; (J-N) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 8.
Juvenile Stage I of Procambarus virginalis. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm.
Figure 9.
Juvenile Stage II of Procambarus virginalis. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D-H) pereiopods 1-5, dorsal or lateral view; (I-M) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 10.
Juvenile Stage II of Procambarus virginalis. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) flagellum of antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bar: 0.25 mm.
Figure 11.
Juvenile Stage III of Procambarus virginalis. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D-H) pereiopods 1-5, dorsal or lateral view; (I-M) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 12.
Juvenile Stage III of Procambarus virginalis. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) flagellum of antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm.
Figure 13.
Juvenile Stage I of Cambaroides japonicus. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan and telson thread, dorsal view; (D) posterior edge of the telson and telson thread; (E-I), pereiopods 1-5, dorsal or lateral view; (J-N) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 14.
Juvenile Stage I of Cambaroides japonicus. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view, (J) pleopod. Scale bars: 0.25 mm.
Figure 15.
Juvenile Stage II of Cambaroides japonicus. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D) posterior edge of tail fan; (E-I) pereiopods 1-5, dorsal or lateral view; (J-N) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 16.
Juvenile Stage II of Cambaroides japonicus. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm.
Figure 17.
Juvenile Stage III of Cambaroides japonicus. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D) posterior edge of tail fan; (E-I) pereiopods 1-5, dorsal or lateral view; (J-N) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.5 mm in A-C, 0.25 mm in D-M.
Figure 18.
Juvenile Stage III of Cambaroides japonicus. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm.
Figure 19.
Juvenile Stage I of Cherax destructor. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D) posterior edge of tail fan; (E-I) pereiopods 1-5, dorsal or lateral view; (J-N) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 20.
Juvenile Stage I of Cherax destructor. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm.
Figure 21.
Juvenile Stage II of Cherax destructor. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D) posterior edge of tail fan; (E-I) pereiopods 1-5, dorsal or lateral view; (J-N) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 22.
Juvenile Stage II of Cherax destructor. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm.
Figure 23.
Juvenile Stage III of Cherax destructor. (A) Habitus, lateral view; (B) eyes and rostrum, dorsal view; (C) tail fan, dorsal view; (D-H) pereiopods 1-5, dorsal or lateral view; (I-M) distal segments of pereiopods 1-5, dorsal or lateral view. Scale bars: 0.25 mm.
Figure 24.
Juvenile Stage III of Cherax destructor. (A) Antenna 1, dorsal view; (B) antennal scale, dorsal view; (C) flagellum of antenna 2, dorsal view; (D) mandible, view of inside of mouth; (E) maxilla 1, oral view; (F) maxilla 2, oral view; (G) first maxilliped, oral view; (H) second maxilliped, oral view; (I) third maxilliped, oral view; (J) pleopod. Scale bars: 0.25 mm.
Figure 25.
Phylogenetic tree of freshwater crayfish, Astacidea, with diagnostic morphological features in postembryonic stages. Basic phylogenetic tree constructed using recent molecular studies (
Crandall et al., 2000Crandall, K.A. Harris, D.J. and Fetzner, J.W. 2000. The monophyletic origin of freshwater crayfish estimated from nuclear and mitochondrial DNA sequences. Proceedings of the Royal Society B, 267(1453): 1679-1686.;
Porter et al., 2005Porter, M.L.; Pérez-Losada, M. and Crandall, K.A. 2005. Model-based multi-locus estimation of decapod phylogeny and divergence times. Molecular Phylogenetics and Evolution, 37: 355-369.;
Bracken et al., 2009Bracken, H.D.; Toon, A.; Felder, D.L.; Martin, J.W.; Finley, M.; Rasmussen, J.; Palero, F. and Crandall, K.A. 2009. The Decapod tree of life: compiling the data and moving toward a consensus of Decapod evolution. Arthropod Systematics & Phylogeny, 67: 99-116.;
Breinholt et al., 2009Breinholt, J.; Pérez-Losada, M. and Crandall, K.A. 2009. The timing of the diversification of the freshwater crayfishes. p. 343-355. In: J.W. Martin; K.A. Crandall and D.L. Felder (eds), Decapod Crustacean Phylogenetics. Crustacean Issues 18, Boca Raton, FL, CRC Press.;
Pérez-Losada et al., 2010Pérez-Losada, A.M.; Schweitzer, C.E.; Feldmann, R.M.; Carlson, M. and Crandall, K.A. 2010. Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules. Journal of Biogeography, 37(12): 2275e2290.;
Toon et al., 2010Toon, A.; Pérez-Losada, M.; Schweitzer, C.E.; Feldmann, R.M.; Carlson, M. and Crandall, K.A. 2010. Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules. Journal of Biogeography, 37: 2275-2290. ;
Stern and Crandall, 2016Stern, D. and Crandall, K.A. 2016. Phylogenetic estimate of the freshwater crayfish (Decapoda: Astacidea) using morphology and molecules. p. 298-309. In: T. Kawai; Z. Faulkes and G. Scholtz (eds), Freshwater Crayfish: a Global Overview. Boca Raton, FL, CRC Press .;
Crandall and De Grave, 2017Crandall, K.A. and De Grave, S. 2017. An updated classification of the freshwater crayfishes (Decapoda: Astacidea) of the world, with a complete species list. Journal of Crustacean Biology, 37: 615-653.;
Grandjean et al., 2017Grandjean, F.; Mun, H.T.; Han, M.G.; Yin, P.L.; Kawai, T.; Distefano, R.J.; Martin, B.; Roles, A.J. and Austin, C.M. 2017. Rapid recovery of nuclear and mitochondrial genes by genome skimming from Northern Hemisphere freshwater crayfish, Zoologica Scripta, 46: 718-728.;
Wolfe et al., 2019Wolfe, J.M.; Breinholt, J.W.; Crandall, K.A.; Lemmon, A.R.; Lemmon, E.M.; Timm, L.E.; Siddall, M.E. and Bracken-Grissom, H.D. 2019. A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings of the Royal Society B: Biological Sciences, 286: 20190079. https://doi.org/10.1098/rspb.2019.0079.
https://doi.org/10.1098/rspb.2019.0079...
). The bars on the tree represent the juvenile morphology as given in Tab. 1; a solid bar means presence and an open vertical bar absence. Numbers on the bar: 1: compound sessile eye in Stage I and developed eye stalk in Stage II; globular carapace in Stage I lost in Stage II; teeth and processes absent in Stage I but present in Stage II; setae absent on pereiopod in Stage I and long setae present in Stage II; 2: setae absent on antenna 1 and 2 with antennal scale in Stage I, and seta present in Stage II; no setae at tip of the exopodite of the maxilliped in Stage I; the plumose setae appear in Stage II; curved hook on terminal end of the pereiopod absent in Stage II; telson with plumose setae along cephalic margin; 3: maxilla 2 with 1 or 3 long setae in Stage II; 4: maxilla 2 with 2 long setae in Stage II; 5: maxilla 2 with 1 long setae in Stage II; 6: maxilla 2 with no long setae in Stage II.
Figure 26.
Estimate of the evolution of maternal care and accompanying postembryonic development in freshwater crayfish (Astacidea) based on juvenile morphology. The tree is based on other recent molecular phylogenetic trees (
Crandall et al., 2000Crandall, K.A. Harris, D.J. and Fetzner, J.W. 2000. The monophyletic origin of freshwater crayfish estimated from nuclear and mitochondrial DNA sequences. Proceedings of the Royal Society B, 267(1453): 1679-1686.;
Porter et al., 2005Porter, M.L.; Pérez-Losada, M. and Crandall, K.A. 2005. Model-based multi-locus estimation of decapod phylogeny and divergence times. Molecular Phylogenetics and Evolution, 37: 355-369.;
Bracken et al., 2009Bracken, H.D.; Toon, A.; Felder, D.L.; Martin, J.W.; Finley, M.; Rasmussen, J.; Palero, F. and Crandall, K.A. 2009. The Decapod tree of life: compiling the data and moving toward a consensus of Decapod evolution. Arthropod Systematics & Phylogeny, 67: 99-116.;
Breinholt et al., 2009Breinholt, J.; Pérez-Losada, M. and Crandall, K.A. 2009. The timing of the diversification of the freshwater crayfishes. p. 343-355. In: J.W. Martin; K.A. Crandall and D.L. Felder (eds), Decapod Crustacean Phylogenetics. Crustacean Issues 18, Boca Raton, FL, CRC Press.;
Pérez-Losada et al., 2010Pérez-Losada, A.M.; Schweitzer, C.E.; Feldmann, R.M.; Carlson, M. and Crandall, K.A. 2010. Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules. Journal of Biogeography, 37(12): 2275e2290.;
Stern and Crandall, 2016Stern, D. and Crandall, K.A. 2016. Phylogenetic estimate of the freshwater crayfish (Decapoda: Astacidea) using morphology and molecules. p. 298-309. In: T. Kawai; Z. Faulkes and G. Scholtz (eds), Freshwater Crayfish: a Global Overview. Boca Raton, FL, CRC Press .; Crandall and De Grave, 2017;
Grandjean et al., 2017Grandjean, F.; Mun, H.T.; Han, M.G.; Yin, P.L.; Kawai, T.; Distefano, R.J.; Martin, B.; Roles, A.J. and Austin, C.M. 2017. Rapid recovery of nuclear and mitochondrial genes by genome skimming from Northern Hemisphere freshwater crayfish, Zoologica Scripta, 46: 718-728.) and molecular clocks (
Toon et al., 2010Toon, A.; Pérez-Losada, M.; Schweitzer, C.E.; Feldmann, R.M.; Carlson, M. and Crandall, K.A. 2010. Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules. Journal of Biogeography, 37: 2275-2290. ;
Wolfe et al., 2019Wolfe, J.M.; Breinholt, J.W.; Crandall, K.A.; Lemmon, A.R.; Lemmon, E.M.; Timm, L.E.; Siddall, M.E. and Bracken-Grissom, H.D. 2019. A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings of the Royal Society B: Biological Sciences, 286: 20190079. https://doi.org/10.1098/rspb.2019.0079.
https://doi.org/10.1098/rspb.2019.0079...
). Juvenile morphology relating to maternal care (
Tab. 5) is shown on the tree. The geological ages and formation of the continents is based on
Owen (1976Owen, H.G. 1976: Continental displacement and expansion of the earth during the Mesozoic and Cenozoic. Philosophical Transactions of the Royal Society A, 281(1303): 223-291.) and
Dercourt et al. (1993Dercourt, J.; Ricou, L. and Vrielynck, B. 1993. Atlas Téthys Palaeoenvironmental Maps: Explanatory Notes. Paris, France, Gauthier-Villars, 307p., 14 maps, 1 pl.). Solid bars indicate presence and blank bars absence. Numbers on the bar: 1: compound sessile eye in Stage I and developed eye stalk in Stage II; 2: globular carapace in Stage I lost in Stage II; 3: teeth and processes on mandible absent in Stage I but present in Stage II; 4: setae absent on pereiopod in Stage I but long setae present in Stage II; 5: no setae on antenna 1-2, with antennal scale in Stage I but seta present in Stage II; 6: no setae at tip of exopodite of maxilliped in Stage I, the plumose setae appear in Stage II; 7: curved hook at tip of the pereiopod present in Stage II; 8: plumose setae on telson in Stage II; 9: setae on maxilla 1 in Stage I; 10: no plumose setae on pleopod in Stage I but present in Stage II. Bold text (Cambaridae and Parastacidae): independent after Stage III; plain text (Astacidae and Cambaroididae): independent after Stage II (
Andrews, 1907Andrews, E.A. 1907b. The attached young of the crayfish Cambarus clarkii and Cambarus diogenes. American Midland Naturalist, 41: 253-271, 2 pls.a;
Albrecht, 1982Albrecht, H. 1982. Das System der europäischen Flußkrebse (Decapoda, Astacidae): Vorschlag und Begrün dung. Mitteilungen des Hamburgischen Zoologischen Museum und Institut, 79: 187-210.;
Scholtz, 1995Scholtz, G. 1995. The attachment of the young in the New Zealand freshwater crayfish Paranephrops zealandicus (White, 1847) (Decapoda, Astacida, Parastacidae). New Zealand Natural Science, 22: 81-89.;
Scholtz and Kawai, 2002Scholtz, G. and Kawai, T. 2002. Aspects of embryonic and postembryonic development of the Japanese freshwater crayfish Cambaroides japonicus (Crustacea, Decapoda) including a hypothesis on the evolutional maternal care in the Astacida. Acta Zoologica, 83: 203-12.).
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