Larval development of Mooreonuphis stigmatis (Treadwell, 1922) (Polychaeta: Onuphidae) from the north-east Pacific

Figures & data

Table I. Summary of different developmental patterns in onuphids (pattern types sensu Paxton (1993) with modifications: I – brooding in the parental tube with direct development; II – viviparity; III – brooding in masses or sacs attached to parental tube; IV – free-spawning with planktonic larvae).

Species	Egg size (µm)	Brood size	Brood care	Type of larva	Type of development	Habitat	Location	Reference
Americonuphis magna (Andrews, 1891)	?	?	Eggs in gelatinous strings	Planktonic, lecithotrophic development	III	Intertidal	North Carolina, USA	Hartman 1945
Brevibrachium maculatum (Estcourt, 1966) as Rhamphobrachium sp.	300–500	10–20	Larvae develop in brood chambers inside the parental tube, egg laying is staggered, 1- to 28-chaetiger larvae found inside one parental tube	Direct development	I	Intertidal	Southern Australia	Smith & Jensz 1968; Paxton 1986
Diopatra albimandibulata Paxton, 1993	?	250	Larvae develop in globular small egg sacs attached to the distal end of parental tube, larvae enclosed by a mucous envelope	Direct development	III	5–18.5 m	Australia, Queensland, Bowen	Paxton 1993
Diopatra? brevicirris Grube, 1856 as D. cuprea Bosc, 1802	?	?	11-chaetiger larvae in the parental tube	Direct development	I		Madeira	Monro 1924; Paxton 1993
Diopatra cuprea Bosc, 1802	220	?	Eggs released in jelly mass which readily dissolved in seawater, larvae reared in the laboratory for 21 days	Planktonic, lecithotrophic development	IV		North East of USA	Allen 1959; Paxton 1993
Diopatra gigova Paxton, 1993	1400	9	Eggs develop in the thin mucous sac on the dorsal side of the worm inside parental tube attached to the tube 1.5 cm from its distal end	? Direct development	I	Intertidal	Western Australia, Dampier Archipelago	Paxton 1993
Diopatra lilliputiana Paxton, 1993	400–900	6–40	Eggs and larvae develop in mucous sac inside parental tube at least to the 15-chaetiger stage	Direct development	I	Intertidal	Western Australia, Broome, Enderby Island, Dampier Archipelago, Noth West cape	Paxton 1993
Diopatra maculata Paxton, 1993	350	1000's	Larvae develop inside mucous matrix which covers the outer distal part of the tube	Direct development	III	Intertidal	Western Australia, Nothern Territory, Queensland	Paxton 1993
Diopatra marocensis Paxton et al., 1995	600	20–100	Larvae develop inside the female tube	Direct development	I	16–40 m	Morocco, Atlantic coast, Sidi Boulbra, fine sand in Abra abra community	Fadlaoui et al. 1995
Diopatra neapolitana Delle Chiaje, 1841	?	?	Eggs develop in the gelatinous mass attached to the distal end of the parental tube	?	III	Intertidal	Italy	Lo Bianco 1898
Diopatra neapolitana Delle Chiaje, 1841	?	?	Artificial fertilization, larvae reared in the laboratory	Planktonic, lecithotrophic development	IV		Arachon, France	Cazaux 1972
Diopatra sp. as D. amboinensis Audouin & Milne-Edwards, 1833	?	?	Up to 6-chaetiger larvae inside the parental tube	Direct development	I		Indonesia as Java	Lieber 1931, cited Paxton 1993
Diopatra sp. as D. sp.nov.	‘Gigantic’	?	Larvae found in the parental tube	Direct development	I		Indonesia as Sumatra	Lieber 1931, cited Paxton 1993
Diopatra sugokai Izuka, 1912 as D. neapolitana Delle Chiaje, 1841	200	?	Eggs in jelly mass attached to the tube opening, artificial fertilization, larvae reared in the laboratory	Planktonic, lecithotrophic development	IV	Intertidal, sandy flat	Japan	Choe 1960
Diopatra tuberculantennata Budaeva & Fauchald, 2008	?	5	Five 28-chaetiger larvae were found in the parental tube	Direct development	I	0–2 m	Caribbean Sea, Belize, west of Dangriga	Budaeva & Fauchald 2008
Diopatra sp.	?	?	6 juveniles (8–15 mm) in mud cocoon	Direct development	? I		South Africa	Day 1960
Diopatra variabilis Southern, 1921	600	30–50	Eggs and larvae of various stages develop in jelly membrane inside parental tube, reared to 15-chaetiger stage in the laboratory	Direct development	I	Shallow brackish water	India	Krishnan 1936; Krishnamoorthi 1963
Fauchaldonuphis paradoxa (Quatrefages, 1866)	Not less than 1200	?2	Two 90-chaetiger larvae about 15 mm in length inside parental tube	Direct development	I	20–72 m	Gulf of Mastatan, Mozambique Channel	Paxton 2005
Hyalinoecia araucana Carrasco, 1983	?	45–69	3–13-chaetiger larvae found inside parental tubes	Direct development	I	600 m	Central Chile, off Lebu	Carrasco 1983
Hyalinoecia bermudensis (Hartman, 1965) as Paronuphis bermudensis	?	3	The anterior half of the body has large ova, and the posterior half contains two well developed embryos	Direct development	II	700–2500 m	Off Bermuda	Hartman 1965
Kinbergonuphis dorsalis (Ehlers, 1897)	?	45	Larvae at least up to 25-chaetiger stage develop in a translucent thin-walled folded bag inside parental tube	Direct development	I	Intertidal	Argentina, Patagonia, Puerto Madryn	Orensanz 1990
Kinbergonuphis notialis (Monro, 1930) as Nothria notialis	?	12–28	Female tube has 6–14 partially isolated external lateral capsules with two ova or developing (up to 24 chaetigers) larvae in each capsule	Direct development	I	210–2900 m	Antarctic	Hartman 1967a, b
Kinbergonuphis simoni (Santos et al., 1982)	370	20–30	Larvae develop up to 20-chaetiger stage in cylindrical membranous chambers inside maternal tube	Direct development	I	Shallow water sandy bay, sorted sediment	Tampa Bay, Florida, USA	Hsieh & Simon 1987
Leptoecia vivipara (Averincev, 1972)	?	7	7–13-chaetiger larvae develop in the body cavity	Direct development	II	437–1180 m	Off Enderby Land and Christensen Coast, Antarctic	Averincev 1972 (as Paronuphis abyssorum); Orensanz 1990
Mooreonuphis jonesi Fauchald, 1982	175–190	50–60	Numerous larvae representing different stages found in a single distinct ten-layerd ovoid chamber inside parental tube	Direct development	I	Shallow subtidal, sandy environments	Off Bermuda	Fauchald 1982b
Mooreonuphis stigmatis (Treadwell, 1922)	400	5–16	Up to 18-chaetiger larvae were found in chambers with very thin walls inside parental tube	Direct development	I	Intertidal	San Juan Island, False Bay, Washington, USA	Present study
Notonuphis antarctica (Monro, 1930) as Paronuphis antarcticus	1170	24	Larvae develop at least up to 24-chaetiger stage inside parental tube	Direct development	I	220–1153 m	Antarctic	Hartman 1967a, b; Averincev 1972
Onuphis elegans (Johnson, 1901) as Nothria elegans	230–235	?	Larvae reared in the laboratory from fertilized egg to 60-chaetiger juvenile	Planktonic, lecithotrophic development	IV	Intertidal –23 m	Dillon Beach, California, USA	Blake 1975

Figure 1.  Early development of Mooreonuphis stigmatis. (A–D) 5-chaetiger larva: (A) lateral view; (B) frontal view; (C,D) fragment of akrotroch. (E,F) 7-chaetiger larva: (E) lateral view, (F) ventral view. (G,H) 11-chaetiger larva: (G) lateral view; (H) ventral view. ak, akrotroch; dac, dorsal anal cirrus; fl, frontal lip; la, lateral antenna; ma, median antenna; me, metatroch; p, palp; pg, pigidium; pr, prototroch; pro, prostomium; vac, ventral anal cirrus.

Figure 2.  16-chaetiger larva of Mooreonuphis stigmatis: (A) dorsal view; (B) prostomium, dorsal view; (C) lateral view; (D) median antenna; (E) parapodium of the third chaetiger; (F) dorsal cirrus of the third parapodium. cil, ciliae; dac, dorsal anal cirrus; dc, dorsal cirrus; la, lateral antenna; ma, median antenna; n, notochaeta, ng, nuchal groove, p, palp; pc, peristomial cirrus; pso, parapodial sensory organ; vac, ventral anal cirrus; vc, ventral cirrus.

Figure 3.  118-chaetiger juvenile of Mooreonuphis stigmatis: (A) lateral view; (B) parapodial sensory organ of the fifth parapodium; (C) anterior end, dorsal view; (D) anterior end, ventral view. dc, dorsal cirus; fl, frontal lip; la, lateral antenna; ll, lower lip; ma, median antenna; ng, nuchal groove; p, palp; pc, peristomial cirrus; psl, postchaetal lobe; pso, parapodial sensory organ; ul, upper lip; vc, ventral cirrus.

Table II. Different types of provisional larval (lowercase) and permanent adult (uppercase) chaetae in comparison with terminology proposed by other authors.

Species			Mooreonuphis stigmatis		Kinbergonuphis simoni		Onuphis elegans		Hyalinoecia araucana		Hirsutonuphis macrocerata
Type of chaeta			Name	Code	Name	Code	Name	Code	Name	Code	Name	Code
Notopodia			Notopodial capillary chaeta	n	Simple capillary chaeta	n	Simple capillary chaeta	d	Larval simple capillary chaeta	L	–	–
Neuropodia	Falcigers	Anterior modified parapodia	Anterior pseudocompound bidentate falciger	a	Pseudocompound bidentate and unidentate falcigers	a, b	–	–	–	–	–	–
			Anterior early compound bidentate hooded falciger	b^1	–	–	–	–	–	–	–	–
			Anterior pseudocompound bidentate hooded falciger	b^2	–	–	–	–	–	–	–	–
			Anterior compound bidentate hooded falciger	b^3	Compound bidentate hooded hooks	e	Larval compound falcigers limited to first two chaetigers	a^1, a^2	Larval compound falcigers	G, H, I	Composite bidentate falciger	–
			Anterior pseudocompound tridentate hooded falciger	B	Pseudocompound bidentate hooded hook	B	Adult compound falcigers limited to first three chaetigers	a^3	–	–	–	–
		Transitional parapodia	Transitional compound bidentate falciger	c	Compound bidentate falciger	c	Larval compound falcigers limited to chaetigers 2-6	b	Larval compound falciger	?F	Compound median and ventral anterior provisional subacicular hooks	m.a.p.SAH v.a.p.SAH
	Hooks	Posterior parapodia with subacicular hooks (SAH)	Posterior compound bidentate subacicular hooded hook	d	Compound bidentate hooded hook	d	Larval compound falciger limited to chaetiger 5 and successive chaetigers	c	Larval compound falciger	J, K	Posterior provisional subacicular hook	p.p.SAH
			Posterior simple subacicular bidentate hooded hook	D	Subacicular bidentate hooded hook	D	Subacicular hook	f	–	–	Permanent subacicular hook	perm.SAH
	Other chaetae		Large median tridentate hooded hook	G	–	–	–	–	–	–	–	–
			Acicular chaeta	A	Acicular chaeta	A	–	–	–	–	–	–
			Simple limbate chaeta	C	Simple limbate chaeta	C	Simple unilimbate capillary chaeta	e	Larval bilimbate capillary chaeta	M	–	–
			Pectinate chaeta	E	–	–	–	–	Larval comb chaeta	N	–	–
			compound spiniger	F	–	–	–	–	–	–	–	–
Reference			Present study		Hsieh & Simon 1987		Blake 1975		Carrasco 1983		Paxton 1996

Figure 4.  Chaetae found on different larvae of Mooreonuphis stigmatis: (A) anterior early compound bidentate hooded falciger – b ¹ ; (B) anterior pseudocompound bidentate falciger – a; (C) anterior pseudocompound bidentate hooded falciger – b ² ; (D) anterior pseudocompound tridentate hooded falciger – B; (E) anterior compound bidentate hooded falciger – b ³ and transitional compound bidentate falciger – c; (F,G) transitional compound bidentate falciger – c; (H) posterior compound bidentate subacicular hooded hook – d; (I) posterior compound bidentate subacicular hooded hook – d, simple limbate chaeta – C and posterior simple subacicular bidentate hooded hook – D. Lowercase letters represent provisional larval chaetae, uppercase letters represent permanent adult chaetae.

Figure 5.  Chaetae present on adults of Mooreonuphis stigmatis: (A) pseudocompound tridentate hooded falciger – B; (B) compound spiniger – F; (C) large median tridentate hooded hook – D; (D) pectinate chaeta – E.

Table III. The distribution of different chaetae during the development of Mooreonuphis stigmatis. Lowercase letters represent provisional larval chaetae (a – anterior pseudocompound bidentate falciger, b ¹ – anterior early compound bidentate hooded falciger, b ² – anterior pseudocompound bidentate hooded falciger, b ³ – anterior compound bidentate hooded falciger, c – transitional compound bidentate falciger, d – posterior compound bidentate subacicular hooded hook, n – notopodial capillary chaeta). Uppercase letters represent permanent adult chaetae (A – acicula, B – anterior pseudocompound tridentate hooded falciger, C – simple limbate chaeta, D – posterior simple subacicular bidentate hooded hook, E – pectinate chaeta). Numbers represent the number of each chaeta on the parapodium of corresponding chaetiger.

	Chaetiger
Stage	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22
5-chaetiger	1b^1	1b^1	1b^1	1b^1
	1A	1A	1A	1A	1A
7-chaetiger	1n	1n	1n
	2a	3c	3c	3c	2c	2c
	1A	1A	1A	1A	1A	1A	1A
10-chaetiger	1n	1n	1n	1n
	2a	3c	3c	3c	3c	3c	1c
	3b^2						2d	1d	1d
	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A
11-chaetiger	1n	1n	1n	1n	1n
	2a	3c	3c	3c	3c	3c
	3b^2						3d	3d	2d
	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A
16-chaetiger	1n	1n	1n	1n	1n
	2a	3c	3c	3c	3c	3c	3c	2c
	3B	1b^3						1d	2d	2d	3d	2d	2d	2d
		2C	2C	2C	1C	1C	1C	1C	2C	2C	2C	2C	1C
									1D
	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A
22-chaetiger	1n	1n	1n
	4b^3	4b^3	3c	3c	3c	3c	3c	3c
		3B							2d	1d	1d	1d	1d	1d	1d	1d	1d	2d	2d	2d	1d
			2C	3C	3C	2C	2C	2C	2C	2C	2C	2C	2C	2C	2C	2C	2C	1C	1C
									1D	1D	1D	1D	1D	1D	1D	1D	1D
												1E	1E	1E	1E	1E	1E
	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A	1A

Table IV. A comparison of the subacicular hooks (SAH) development among ten species of onuphids.

Species	Stage when first SAH appear	No. chaetiger with the first SAH in juveniles	No. chaetiger with the first SAH in adults	Reference
Diopatra lilliputiana	15-chaetiger	6	14 (10–12)	Paxton 1993
Diopatra marocensis	12-chaetiger	9	13–15	Fadlaoui et al. 1995
Fauchaldonuphis paradoxa	90-chaetiger	9	15–17	Paxton 2005
Hirsutonuphis armillata	?	11	11–12	Paxton 1996
Hirsutonuphis intermedia	?	10–11	13–15	Paxton 1996
Hirsutonuphis macrocerata	?	10–11	15–19	Paxton 1996
Kinbergonuphis simoni	17-chaetiger	9	17	Hsieh & Simon 1987
Mooreonuphis stigmatis	16-chaetiger	9	16	Present study
Onuphis elegans	14-chaetiger	8	9	Blake 1975
Rhamphobrachium ehlersi	?	7–8	15–16	Paxton 1986

Table V. A comparison of the development of branchiae among five species of onuphids.

Species	Stage when first branchiae appear	No. chaetiger with first branchiae in larvae	Range of chaetigers with branchiae in adults	Reference
Diopatra marocensis	18–20-chaetiger	5	4(5)–19(43)	Fadlaoui et al. 1995
Kinbergonuphis simoni	36-chaetiger	9–10	6(9)–20	Hsieh & Simon 1987
Mooreonuphis jonesi	10-chaetiger	6–8	29–85	Fauchald 1982b
Mooreonuphis stigmatis	Later than 24-chaetiger	?	16(20)– posterior	Present study
Onuphis elegans	9-chaetiger	6–7	1– posterior	Blake 1975

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