Diversity of hippoidean crabs - considering ontogeny, quantifiable morphology, and phenotypic plasticity

Braig, Florian; Zuluaga, Victor Posada; Haug, Carolin; Haug, Joachim T.

doi:10.1590/2358-2936e2021027

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Original Article • Nauplius 29 • 2021 • https://doi.org/10.1590/2358-2936e2021027 copy

Diversity of hippoidean crabs - considering ontogeny, quantifiable morphology, and phenotypic plasticity

Authorship SCIMAGO INSTITUTIONS RANKINGS

Abstract

Representatives of Hippoidea, often called sand crabs or mole crabs, are an ingroup of Anomala. These marine crustaceans inhabit the tropical and subtropical coasts of the world, yet some also appear in temperate climates. Their adults are specialized for digging and living in sandy substrates. Hippoidean zoea-type larvae are planktic and reach large sizes up to a few centimetres. These larvae transform into megalopa larvae, strongly resembling the adult, mediating the transition to the benthic lifestyle of the adult. We reconstructed outlines in dorsal view of over 80 shields of hippoideans, including representatives of Blepharipodidae (sister group to all others), Albuneidae, and Hippidae and including adults, megalopa-type, and zoea-type larvae from all three ingroups. We conducted a morphological analysis on this data using an elliptic Fourier transformation and principal component analysis. We used the results of the analysis to discuss the life history of hippoideans and the special function of megalopae, which often lack emphasis in current research. Early stage zoea larvae, megalopae, and adults show a linear gradient in their morphological development according to our analysis. However, late stage zoea larvae deviate from this pattern, possibly due to their specialization to a long-lasting planktic life. Lastly, we discuss the influence of phenotypic plasticity in hippoidean zoea larvae.

Keywords:
Eucrustacea; Fourier analysis; Hippoidea; life phases; morphometrics

no.	major group	species group	species	size class	lab/wild	author	year	Figure	accession number	museum	geographic information	cruise	further info
1	Hippidae	Emerita	analoga	adult		*Uribe et al.*Uribe, R.A.; Rubio, J.R.; Carbajal, P.E. and Berrú, P.P. 2013. Invertebrados marinos bentónicos del litoral de la Región Áncash, Perú. Boletín del Instituto del Mar del Perú, 28: 136-293.	2013	Emerita analoga (Stimpson, 1857)
2	Hippidae	Emerita	analoga	late zoea	Lab	PulsPuls, A.L. 2001. Arthropoda: Decapoda. p. 179-250. In: A. Shanks (ed), An identification guide to the larval marine invertebrates of the Pacific Northwest. Corvallis, Oregon State University Press.	2001	fig. 18C
3	Hippidae	Emerita	brasiliensis	adult		ScelzoScelzo, M.A. 2004. Cangrejos Anomura: Ermitaños y chinches de la arena. p. 213-218. In: E.E. Boschi and M.B. Cousseau (eds), La vida entre mareas: vegetales y animales de las costas de Mar del Plata, Argentina. Mar de Plata, Instituto Nacional de Investigación y Desarrollo Pesquero.	2004	fig. 1B
4	Hippidae	Emerita	emiritus	adult		GomalanonGomalanon, P. 2016. Species and distribution of sand crabs (Crustacea: Hippoidea) in Chalathat Beach, Songkhla Province. Songkhla, Prince of Songkla University, 100p., Avilable at , Avilable at https://kb.psu.ac.th/psukb/bitstream/2016/11390/1/416972.pdf . Accessed on 15 February 2020. [In Thai with English abstract]. https://kb.psu.ac.th/psukb/bitstream/201...	2016	Fig. 6
5	Hippidae	Emerita	holthuisi	adult		SankolliSankolli, K.N. 1965. On a new species of Emerita (Decapoda, Anomura) from India, with a note on Emerita emeritus (L.). Crustaceana, 8: 48-54.	1965	fig. 1A
6	Hippidae	Emerita	holthuisi	megalopa		*Harvey et al.*Harvey, A.; Boyko, C.B.; McLaughlin, P. and Martin, J.W. 2014. Anomura. p. 283-287. In: J.W. Martin, J. Olesen and J.T. Høeg (eds), Atlas of crustacean larvae. Baltimore, The Johns Hopkins University Press.	2014	fig. 53.6B
7	Hippidae	Emerita	holthuisi	early zoea	Lab	*Harvey et al.*Harvey, A.; Boyko, C.B.; McLaughlin, P. and Martin, J.W. 2014. Anomura. p. 283-287. In: J.W. Martin, J. Olesen and J.T. Høeg (eds), Atlas of crustacean larvae. Baltimore, The Johns Hopkins University Press.	2014	fig. 53.1C
8	Hippidae	Emerita	holthuisi	late zoea	Lab	Siddiqi and GhorySiddiqi, F.A. and Ghory, F.S. 2006. Complete larval development of Emerita holthuisi Sankolli, 1965 (Crustacea: Decapoda: Hippidae) reared in the laboratory. Turkish Journal of Zoology, 30: 121-135.	2006	fig. 2A
9	Hippidae	Emerita	holthuisi	late zoea	Lab	*Harvey et al.*Harvey, A.; Boyko, C.B.; McLaughlin, P. and Martin, J.W. 2014. Anomura. p. 283-287. In: J.W. Martin, J. Olesen and J.T. Høeg (eds), Atlas of crustacean larvae. Baltimore, The Johns Hopkins University Press.	2014	fig. 53.1D
10	Hippidae	Emerita	holthuisi	late zoea	Lab	Siddiqi and GhorySiddiqi, F.A. and Ghory, F.S. 2006. Complete larval development of Emerita holthuisi Sankolli, 1965 (Crustacea: Decapoda: Hippidae) reared in the laboratory. Turkish Journal of Zoology, 30: 121-135.	2006	fig. 4A
11	Hippidae	Emerita	holthuisi	late zoea	Lab	Siddiqi and GhorySiddiqi, F.A. and Ghory, F.S. 2006. Complete larval development of Emerita holthuisi Sankolli, 1965 (Crustacea: Decapoda: Hippidae) reared in the laboratory. Turkish Journal of Zoology, 30: 121-135.	2006	fig. 5A
12	Hippidae	Emerita	holthuisi	late zoea	Lab	Siddiqi and GhorySiddiqi, F.A. and Ghory, F.S. 2006. Complete larval development of Emerita holthuisi Sankolli, 1965 (Crustacea: Decapoda: Hippidae) reared in the laboratory. Turkish Journal of Zoology, 30: 121-135.	2006	fig. 6A
13	Hippidae	Emerita	portoricensis	adult		SchmittSchmitt, W.L. 1935. Crustacea Macrura and Anomura of Porto Rico and the Virgin Islands. Scientific Survey of Porto Rico and the Virgin Islands, 15: 125- 227.	1935	fig. 72B
14	Hippidae	Emerita	rathbunae	megalopa		KnightKnight, M.D. 1967. The larval development of the sand crab Emerita rathbunae Schmitt (Decapoda, Hippidae). Pacific Science, 21: 58-76.	1967	fig. 36
15	Hippidae	Emerita	rathbunae	late zoea	Wild	KnightKnight, M.D. 1967. The larval development of the sand crab Emerita rathbunae Schmitt (Decapoda, Hippidae). Pacific Science, 21: 58-76.	1967	fig. 7
16	Hippidae	Emerita	rathbunae	late zoea	Lab	KnightKnight, M.D. 1967. The larval development of the sand crab Emerita rathbunae Schmitt (Decapoda, Hippidae). Pacific Science, 21: 58-76.	1967	fig. 8
17	Hippidae	Emerita	sp.	megalopa		FonghoyFonghoy, C. 2015. Ontogeny and larval development of sand crab, Emerita sp. (Decapoda: Anomura: Hippidae) reared in laboratory. Bangkok, Chulalongkorn University, Department of Marine Science, Masters Thesis, 88p., Availabe at , Availabe at http://cuir.car.chula.ac.th/handle/123456789/60930 . Accessed on 15 February 2020. [Unpublished] http://cuir.car.chula.ac.th/handle/12345...	2015	fig. 37A
18	Hippidae	Emerita	sp.	early zoea	Lab	FonghoyFonghoy, C. 2015. Ontogeny and larval development of sand crab, Emerita sp. (Decapoda: Anomura: Hippidae) reared in laboratory. Bangkok, Chulalongkorn University, Department of Marine Science, Masters Thesis, 88p., Availabe at , Availabe at http://cuir.car.chula.ac.th/handle/123456789/60930 . Accessed on 15 February 2020. [Unpublished] http://cuir.car.chula.ac.th/handle/12345...	2015	fig. 25A
19	Hippidae	Emerita	sp.	late zoea	Lab	FonghoyFonghoy, C. 2015. Ontogeny and larval development of sand crab, Emerita sp. (Decapoda: Anomura: Hippidae) reared in laboratory. Bangkok, Chulalongkorn University, Department of Marine Science, Masters Thesis, 88p., Availabe at , Availabe at http://cuir.car.chula.ac.th/handle/123456789/60930 . Accessed on 15 February 2020. [Unpublished] http://cuir.car.chula.ac.th/handle/12345...	2015	fig. 27A
20	Hippidae	Emerita	sp.	late zoea	Lab	FonghoyFonghoy, C. 2015. Ontogeny and larval development of sand crab, Emerita sp. (Decapoda: Anomura: Hippidae) reared in laboratory. Bangkok, Chulalongkorn University, Department of Marine Science, Masters Thesis, 88p., Availabe at , Availabe at http://cuir.car.chula.ac.th/handle/123456789/60930 . Accessed on 15 February 2020. [Unpublished] http://cuir.car.chula.ac.th/handle/12345...	2015	fig. 29A
21	Hippidae	Emerita	sp.	late zoea	Lab	FonghoyFonghoy, C. 2015. Ontogeny and larval development of sand crab, Emerita sp. (Decapoda: Anomura: Hippidae) reared in laboratory. Bangkok, Chulalongkorn University, Department of Marine Science, Masters Thesis, 88p., Availabe at , Availabe at http://cuir.car.chula.ac.th/handle/123456789/60930 . Accessed on 15 February 2020. [Unpublished] http://cuir.car.chula.ac.th/handle/12345...	2015	fig. 31A
22	Hippidae	Emerita	sp.	late zoea	Lab	FonghoyFonghoy, C. 2015. Ontogeny and larval development of sand crab, Emerita sp. (Decapoda: Anomura: Hippidae) reared in laboratory. Bangkok, Chulalongkorn University, Department of Marine Science, Masters Thesis, 88p., Availabe at , Availabe at http://cuir.car.chula.ac.th/handle/123456789/60930 . Accessed on 15 February 2020. [Unpublished] http://cuir.car.chula.ac.th/handle/12345...	2015	fig. 33A
23	Hippidae	Emerita	sp.	late zoea	Lab	FonghoyFonghoy, C. 2015. Ontogeny and larval development of sand crab, Emerita sp. (Decapoda: Anomura: Hippidae) reared in laboratory. Bangkok, Chulalongkorn University, Department of Marine Science, Masters Thesis, 88p., Availabe at , Availabe at http://cuir.car.chula.ac.th/handle/123456789/60930 . Accessed on 15 February 2020. [Unpublished] http://cuir.car.chula.ac.th/handle/12345...	2015	fig. 35A
24	Hippidae	Emerita	taiwanensis	adult		HsuehHsueh, P.W. 2015. A new species of Emerita (Decapoda, Anomura, Hippidae) from Taiwan, with a key to species of the genus. Crustaceana, 88: 247-258.	2015	fig. 2A
25	Hippidae	Emerita	talpoida	adult		GomalanonGomalanon, P. 2016. Species and distribution of sand crabs (Crustacea: Hippoidea) in Chalathat Beach, Songkhla Province. Songkhla, Prince of Songkla University, 100p., Avilable at , Avilable at https://kb.psu.ac.th/psukb/bitstream/2016/11390/1/416972.pdf . Accessed on 15 February 2020. [In Thai with English abstract]. https://kb.psu.ac.th/psukb/bitstream/201...	2016	fig. 9
26	Hippidae	Hippa	adactyla	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1G
27	Hippidae	Hippa	granulatus	adult		BorradaileBorradaile, L.A. 1904. Marine crustaceans. XIII. The Hippidea, Thalassinidea and Scyllaridea. p. 750-754. In: J. S. Gardiner (ed), The fauna and geography of the Maldive and Laccadive Archipelagoes 2. Cambridge, Cambridge University Press.	1904	fig. 1A
28	Hippidae	Hippa	indica	adult		*Haig et al.*Haig, J.; Murugan, T. and Nair, N.B. 1986. Hippa indica, a new species of mole crab (Decapoda, Anomura, Hippidae) from the south west coast of India. Crustaceana, 51: 286-292.	1986	fig. 1A
29	Hippidae	Hippa	marmorata	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1H
30	Hippidae	Hippa	ovalis	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1I
31	Hippidae	Hippa	strigillata	adult		MiersMiers, E.J. 1878. Revision of the Hippidea. Zoological Journal of the Linnean Society, 14(76): 312-336.	1878	fig. 3
32	Hippidae	Hippa	truncatifrons	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1J
33	Hippidae	Hippa	truncatifrons	megalopa		Kato and SuzukiKato, T. and Suzuki, H. 1992. Some biological observations of the mole crabs (Hippidae, Anomura, Decapoda, Crustacea) in Sagami Bay and the complete larval development of Hippa truncatifrons (Miers). Reports of the Manazuru Marine Laboratory for Science Education, Faculty of Education, Yokohama University, 8: 77-97. (In Japanese with English abstract)	1992	fig. 7A
34	Hippidae	Mastigochirus	gracilis	adult		MiersMiers, E.J. 1878. Revision of the Hippidea. Zoological Journal of the Linnean Society, 14(76): 312-336.	1878	fig. 7
35	Hippidae	Mastigochirus	quadrilobatus	adult		MiersMiers, E.J. 1878. Revision of the Hippidea. Zoological Journal of the Linnean Society, 14(76): 312-336.	1878	fig. 8
36	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 4A-B	MNHN-IU-2014-5475A	MNHN Paris	3°38'S 9°22'E, west of Gabun	Ombango 1960, c. 12, station 301	leg. 03.05.1960
37	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 4C-D	MNHN-IU-2014-5475B	MNHN Paris	3°38'S 9°22'E, west of Gabun	Ombango 1960, c. 12, station 301	leg. 03.05.1960
38	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 3E-F	MNHN-IU-2014-5524A	MNHN Paris	23°07'S 43°11'W, south of Brazil	Calypso 1961-62, station 108	-
39	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 3G-I	MNHN-IU-2014-5524B	MNHN Paris	23°07'S 43°11'W, south of Brazil	Calypso 1961-62, station 108	-
40	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 3J-L	MNHN-IU-2014-5526	MNHN Paris	24°03'S 46°22'W, south of Brazil	Calypso 1961-62, station 139	-
41	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 3A-D	ZMH-K07448A	CeNak Hamburg	20°S 73°W, west of Chile	-	leg. H. Nissen, 23.04.1907
42	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 4E-H	ZMH-K07448B	CeNak Hamburg	20°S 73°W, west of Chile	-	leg. H. Nissen, 23.04.1907
43	Hippidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 1	ZMH-K16356	CeNak Hamburg	Sansibar	-
44	Hippidae	Unknown	Unknown	zoea	Wild	Rudolf et al.Rudolf, N.R.; Haug, C. and Haug, J.T. 2016. Functional morphology of giant mole crab larvae: a possible case of defensive enrollment. Zoological Letters, 2: 17.	2016	fig. 5	MNHN-IU-2014-5468	MNHN Paris
45	Hippidae	Unknown	Unknown	zoea	Wild	*Rudolf et al.*Rudolf, N.R.; Haug, C. and Haug, J.T. 2016. Functional morphology of giant mole crab larvae: a possible case of defensive enrollment. Zoological Letters, 2: 17.	2016	fig. 5	SMF-Mu_267	Senckenberg Naturmuseum Frankfurt
46	Hippidae	Unknown	Unknown	zoea	Wild	*Rudolf et al.*Rudolf, N.R.; Haug, C. and Haug, J.T. 2016. Functional morphology of giant mole crab larvae: a possible case of defensive enrollment. Zoological Letters, 2: 17.	2016	fig. 5	ZMUC-CRU-8679	NHMD Copenhagen
47	Hippidae	Unknown	Unknown	zoea	Wild	*Rudolf et al.*Rudolf, N.R.; Haug, C. and Haug, J.T. 2016. Functional morphology of giant mole crab larvae: a possible case of defensive enrollment. Zoological Letters, 2: 17.	2016	fig. 5	ZMUC-CRU-8680	NHMD Copenhagen
48	Hippidae	Unknown	Unknown	zoea	Wild	*Rudolf et al.*Rudolf, N.R.; Haug, C. and Haug, J.T. 2016. Functional morphology of giant mole crab larvae: a possible case of defensive enrollment. Zoological Letters, 2: 17.	2016	fig. 5	ZMUC-CRU-8682	NHMD Copenhagen
49	Hippidae	Unknown	Unknown	zoea	Wild	*Rudolf et al.*Rudolf, N.R.; Haug, C. and Haug, J.T. 2016. Functional morphology of giant mole crab larvae: a possible case of defensive enrollment. Zoological Letters, 2: 17.	2016	fig. 5	ZMUC-CRU-8683	NHMD Copenhagen
50	Hippidae	Unknown	Unknown	zoea	Wild	*Rudolf et al.*Rudolf, N.R.; Haug, C. and Haug, J.T. 2016. Functional morphology of giant mole crab larvae: a possible case of defensive enrollment. Zoological Letters, 2: 17.	2016	fig. 5	ZMUC-CRU-8684	NHMD Copenhagen
51	Albuneidae	Albunea	carabus	adult		Abdelsalam and RamadanAbdelsalam, K. and Ramadan, S.E. 2017. First record of two crab species from the Egyptian Mediterranean Sea. Cahiers de Biologie Marine, 58: 17-23.	2017	fig. 2A
52	Albuneidae	Albunea	carabus	early zoea	Wild	SeridjiSeridji, R. 1988. Some planktonic larval stages of Albunea carabus (L., 1758) (Crustacea, Decapoda, Anomura). Journal of Natural History, 22: 1293-1300.	1988	fig. 1A
53	Albuneidae	Albunea	carabus	late zoea	Wild	SeridjiSeridji, R. 1988. Some planktonic larval stages of Albunea carabus (L., 1758) (Crustacea, Decapoda, Anomura). Journal of Natural History, 22: 1293-1300.	1988	fig. 2A
54	Albuneidae	Albunea	carabus	late zoea	Wild	SeridjiSeridji, R. 1988. Some planktonic larval stages of Albunea carabus (L., 1758) (Crustacea, Decapoda, Anomura). Journal of Natural History, 22: 1293-1300.	1988	fig. 3A
55	Albuneidae	Albunea	elioti	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1A
56	Albuneidae	Albunea	occulta	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1B
57	Albuneidae	Albunea	symmysta	adult		*Mashar et al.*Mashar, A.; Wardiatno, Y.; Boer, M.; Butet, N.A.; Farajallah, A. and Ardika, P. U. 2015. First record of Albunea symmysta (Crustacea: Decapoda: Albuneidae) from Sumatra and Java, Indonesia. Aquaculture, Aquarium, Conservation & Legislation, 8: 611-615.	2015	fig. 3A
58	Albuneidae	Austrolepidopa	schmitti	adult		Efford and HaigEfford, I.E. and Haig, J. 1968. Two new genera and three new species of crabs (Decapoda: Anomura: Albuneidae) from Australia. Australian Journal of Zoology, 16: 897-914.	1968	fig. 1
59	Albuneidae	Austrolepidopa	trigonops	adult		Efford and Haig,Efford, I.E. and Haig, J. 1968. Two new genera and three new species of crabs (Decapoda: Anomura: Albuneidae) from Australia. Australian Journal of Zoology, 16: 897-914.	1968	fig. 5
60	Albuneidae	Lepidopa	benedicti	adult		FaulkesFaulkes, Z. 2017. The phenology of sand crabs, Lepidopa benedicti (Decapoda: Albuneidae). Journal of Coastal Research, 33: 1095-1101.	2017	fig. 5B (center)
61	Albuneidae	Lepidopa	benedicti	adult		FaulkesFaulkes, Z. 2017. The phenology of sand crabs, Lepidopa benedicti (Decapoda: Albuneidae). Journal of Coastal Research, 33: 1095-1101.	2017	fig. 5B (middle)
62	Albuneidae	Lepidopa	benedicti	adult		FaulkesFaulkes, Z. 2017. The phenology of sand crabs, Lepidopa benedicti (Decapoda: Albuneidae). Journal of Coastal Research, 33: 1095-1101.	2017	fig. 5B (center)
63	Albuneidae	Lepidopa	benedicti	megalopa		*Harvey et al.*Harvey, A.; Boyko, C.B.; McLaughlin, P. and Martin, J.W. 2014. Anomura. p. 283-287. In: J.W. Martin, J. Olesen and J.T. Høeg (eds), Atlas of crustacean larvae. Baltimore, The Johns Hopkins University Press.	2014	fig. 53.6A
64	Albuneidae	Lepidopa	benedicti	early zoea	Lab	Stuck and Truesdale,Stuck, K.C. and Truesdale, F.M. 1986. Larval and early development of Lepidopa benedicti Schmitt, 1935 (Anomura: Albuneidae) reared in laboratory. Journal of Crustacean Biology, 6: 89-110.	1986	fig. 1B
65	Albuneidae	Lepidopa	benedicti	late zoea	Lab	Stuck and TruesdaleStuck, K.C. and Truesdale, F.M. 1986. Larval and early development of Lepidopa benedicti Schmitt, 1935 (Anomura: Albuneidae) reared in laboratory. Journal of Crustacean Biology, 6: 89-110.	1986	fig. 2A
66	Albuneidae	Lepidopa	benedicti	late zoea	Lab	Stuck and TruesdaleStuck, K.C. and Truesdale, F.M. 1986. Larval and early development of Lepidopa benedicti Schmitt, 1935 (Anomura: Albuneidae) reared in laboratory. Journal of Crustacean Biology, 6: 89-110.	1986	fig. 3A
67	Albuneidae	Lepidopa	benedicti	late zoea	Lab	Stuck and TruesdaleStuck, K.C. and Truesdale, F.M. 1986. Larval and early development of Lepidopa benedicti Schmitt, 1935 (Anomura: Albuneidae) reared in laboratory. Journal of Crustacean Biology, 6: 89-110.	1986	fig. 4B
68	Albuneidae	Lepidopa	websteri	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1C
69	Albuneidae	Paraleucolepidopa	myops	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1D
70	Albuneidae	Paraleucolepidopa	myops	late zoea	Lab	*Harvey et al.*Harvey, A.; Boyko, C.B.; McLaughlin, P. and Martin, J.W. 2014. Anomura. p. 283-287. In: J.W. Martin, J. Olesen and J.T. Høeg (eds), Atlas of crustacean larvae. Baltimore, The Johns Hopkins University Press.	2014	fig. 53.1A
71	Albuneidae	Stemonopa	insignis	adult		Efford and HaigEfford, I.E. and Haig, J. 1968. Two new genera and three new species of crabs (Decapoda: Anomura: Albuneidae) from Australia. Australian Journal of Zoology, 16: 897-914.	1968	fig. 8
72	Albuneidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 2G-H	MNHN-IU-2014-5518	MNHN Paris	-	Calypso 1961-62, station 153	-
73	Albuneidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 2A-C	MNHN-IU-2014-5523	MNHN Paris	08°25'S 34°48'W, east of Brazil	Calypso 1961-62, station 26	-
74	Albuneidae	Unknown	Unknown	zoea	Wild	This paper		Fig. 2D-F	MNHN-IU-2014-5527	MNHN Paris	24°03'S 46°22'W, south of Brazil	Calypso 1961-62, station 139	-
75	Blepharipodidae	Blepharipoda	doelloi	adult		SchmittSchmitt, W.L. 1942. A new species of sand bug, Blepharipodida doelloi, from Argentina. Smithsonian Miscellaneous Collections, 101(18): 1-10.	1942	fig. 1
76	Blepharipodidae	Blepharipoda	doelloi	adult		SchmittSchmitt, W.L. 1942. A new species of sand bug, Blepharipodida doelloi, from Argentina. Smithsonian Miscellaneous Collections, 101(18): 1-10.	1942	fig. 3
77	Blepharipodidae	Blepharipoda	doelloi	megalopa		*Harvey et al.*Harvey, A.; Boyko, C.B.; McLaughlin, P. and Martin, J.W. 2014. Anomura. p. 283-287. In: J.W. Martin, J. Olesen and J.T. Høeg (eds), Atlas of crustacean larvae. Baltimore, The Johns Hopkins University Press.	2014	fig. 53.6C
78	Blepharipodidae	Blepharipoda	liberata	adult		ShenShen, C.J. 1949. Notes on the genera Blepharipoda and Lophomastix of the family Albuneidae (Crustacea Anomura) with description of a new species, B. liberata, from China. Contributions of the Institute of Zoology of the National Academy of Peiping, 5: 153-170.	1949	Plate XIV
79	Blepharipodidae	Blepharipoda	occidentalis	adult		Boyko and McLaughlinBoyko, C.B. and McLaughlin, P.A. 2010. Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part IV - Hippoidea. The Raffles Bulletin of Zoology, 23: 139-151.	2010	fig. 1E
80	Blepharipodidae	Blepharipoda	occidentalis	early zoea	Lab	*Harvey et al.*Harvey, A.; Boyko, C.B.; McLaughlin, P. and Martin, J.W. 2014. Anomura. p. 283-287. In: J.W. Martin, J. Olesen and J.T. Høeg (eds), Atlas of crustacean larvae. Baltimore, The Johns Hopkins University Press.	2014	fig. 53.1E
81	Blepharipodidae	Blepharipoda	occidentalis	late zoea	Lab	Johnson and LewisJohnson, M.W. and Lewis, W.M. 1942. Pelagic larval stages of the sand crabs Emerita analoga (Stimpson), Blepharipoda occidentalis Randall, and Lepidopa myops Stimpson. Biological Bulletin, 83: 67-87.	1942	Plate IV, fig. 1
82	Blepharipodidae	Blepharipoda	spinosa	adult		Milne Edwards and LucasMilne Edwards, H. and Lucas, H. 1841. Description des Crustacés nouveaux ou peu connus conservés dans la collection du muséum d’histoire naturelle. Archives du Muséum national d’Histoire naturelle Paris, 2: 461-483.	1841	Plate XXVIII, fig. 1
83	Blepharipodidae	Lophomastix	japonica	adult		DurufléDuruflé, M. 1889. Description d'une nouvelle espèce du genre Blepharipoda. Bulletin de la Sociéte Philomathique de Paris, 8: 92-95.	1889	Blephacopoda Japonica (Duru.) [sic]
84	Blepharipodidae	Lophomastix	japonica	megalopa		KonishiKonishi, K. 1987. Larval development of the spiny sand crab Lophomastix japonica (Durufle, 1889) (Crustacea, Anomura, Albuneidae) under laboratory conditions. Publications of the Seto Marine Biological Laboratory, 32: 123-139.	1987	fig. 7A

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Florian Braig florianbraig@t-online.de

Morpho-larvas. l.	Immature that differs in its morphology from that of the adult.
Morpho-larva s. str.	Immature that differs in its morphology from that of the adult and possesses structures that get reduced later in ontogeny.
Eco-larva s. l.	Immature that differs significantly in its ecological niche from that of the adult.
Eco-larva s. str.	Immature that differs significantly in its ecological niche from that of the adult and fulfils the specific function of dispersal.
Metamorph-larva	Immature that transforms into non-larval stage by metamorphosis.
Apo-larva	Immature that fulfils at least one of the above criteria and possesses evolutionary new structures for this specific stage.
Plesio-larva	Immature that fulfils at least one of the above criteria and possesses no evolutionary new structures for this specific stage.