Description of the complete larval development of Lysmata amboinensis (De Man) (Decapoda: Lysmatidae) reared under laboratory conditions Author De Sousa, Lígia F. 0000-0003-1934-1836 IPMA - Instituto Português do Mar e Atmosfera, Av. Doutor Alfredo Magalhães Ramalho, 6, 1495 - 165 Algés, Portugal. & ligia. faria. sousa @ gmail. com; https: // orcid. org / 0000 - 0003 - 1934 - 1836 & CIIMAR - Centro Interdisciplinar de Investigação Marinha e Ambiental, Terminal de Cruzeiros do Porto de Leixões, Matosinhos, Portugal ligia.faria.sousa@gmail.com Author Marques, Daniel 0000-0001-8247-8531 MARE - Marine and Environmental Sciences Centre, Polytechnic Institute of Leiria, 2520 - 630 Peniche, Portugal. dfmarques 88 @ gmail. com; https: // orcid. org / 0000 - 0001 - 8247 - 8531 dfmarques88@gmail.com Author Leandro, Sérgio Miguel 0000-0001-5005-3598 MARE - Marine and Environmental Sciences Centre, ESTM, Polytechnic Institute of Leiria, 2520 - 614 Peniche, Portugal. sergio. leandro @ ipleiria. pt; https: // orcid. org / 0000 - 0001 - 5005 - 3598 sergio.leandro@ipleiria.pt Author Santos, Antonina Dos 0000-0002-2238-9315 IPMA - Instituto Português do Mar e Atmosfera, Av. Doutor Alfredo Magalhães Ramalho, 6, 1495 - 165 Algés, Portugal. & antonina @ ipma. pt; https: // orcid. org / 0000 - 0002 - 2238 - 9315 & CIIMAR - Centro Interdisciplinar de Investigação Marinha e Ambiental, Terminal de Cruzeiros do Porto de Leixões, Matosinhos, Portugal antonina@ipma.pt text Zootaxa 2022 2022-02-16 5099 5 501 526 journal article 10.11646/zootaxa.5099.5.1 1175-5326 6110178 F52DBA8A-93C6-4950-997A-C1D7484812E0 Morphological description of larval stages of Lysmata amboinensis ( Fig 1–7 ) First Zoea ( Figure 1A–J ) Size: TL= 2.86± 0.04 mm ; CL= 0.95± 0.02 mm ; RL= 0.30± 0.03 mm ; N = 10 Carapace ( Figure 1A , A’): rostrum slender and pointed, reaching half the size of the antennular peduncle, eyes compound and sessile; carapace smooth with 1 pterigostomial spine, followed by 3–5 denticles along anterior ventral margin. Antennule ( Figure 1B ): peduncle entire, with endopod represented by 1 long plumose setae, and exopod with 1–2 plumose setae and 3 aesthetascs. Antenna ( Figure 1C ): peduncle entire; endopod apically with 1 long plumose setae and 1 short spine; exopod with 5-articles, with 9 plumose setae on inner side and 2 plumose setae on outer side, plus a simple setae on apex. Mandible ( Figure 1D ): asymmetrical, palp absent, left mandible bearing 3 teeth and right mandible with 5 teeth, both with 1 lacinia mobilis , molar process as illustrated. Maxillule ( Figure 1E ): coxopodite with 7 cuspidate setae, basipodite with 6 cuspidate setae; endopod with 2 strong subterminal serrulate setae and 3 strong terminal serrulate setae. Maxilla ( Figure 1F ): coxopodite bilobed with 8–9 (3 serrulate and 5–6 pappose) and 4 pappose setae, respectively; basipodite bilobed with 3 pappose setae and 4 pappose setae, respectively; endopod entire bearing 3+2+4 pappose setae; exopod with 5 marginal plumose setae. First maxilliped ( Figure 1G ): coxopodite with 2+1+2 serrulate setae; basipodite with 12 (4 serrulate and 8 pappose) setae; endopod with 4 articles with 3,1,2, 4–5 pappose setae; exopod entire, with 1+2+1 plumose setae. Second maxilliped ( Figure 1H ): coxopodite unarmed; basipodite with 1+2+2 serrulate setae; endopod with 3 articles with 1–3 (2 simple and 1 plumodenticulate) + 0–1 simple, 0–3 (0–2 simple and 0–1 plumodenticulate), 4+1 (2 simple and 3 plumodenticulate) setae; exopod entire bearing 2+2+2+2+1 plumose setae. Third maxilliped ( Figure 1I ): coxopodite unarmed; basipodite with 2+1 serrulate setae; endopod with 4 articles with 2+1 simple, 0, 1+4 plumodenticulate, 2–3 (0–1 simple and 2 plumodenticulate) setae; exopod entire bearing 2+2+2+2+1 plumose setae. First pereiopod ( Figure 1A ): as bud Second to fifth pereiopods: absent Pleon ( Figure 1A ): 5 somites; 1 pair of dorso-lateral spines on posterior margin of the 5 th somite. Pleopods: absent Uropod: absent Telson ( Figure 1A, 1J ): triangular, indented medially with 7+7 posterior processes; minute spines between setae. Second Zoea ( Figure 2A–F ) Size: TL= 3.16± 0.10 mm ; CL= 0.85± 0.11mm ; RL= 0.38± 0.03 mm ; N = 10 Carapace ( Figure 2A ): rostrum slender and pointed, reaching half the size of the antennular peduncle, eyes compound and stalked; 1 supraorbital spine, 1 pterigostomial spine, and 3–6 denticles along ventral margin. Antennule ( Figure 2A ): peduncle with 2 articles, distal article bearing 2 plumose setae distally; endopod represented by 1 long plumose setae; exopod with 1 simple seta and 4 long aesthetascs. Antenna ( Figure 2A, 2B ): peduncle entire; endopod represented by 1 long plumose setae; exopod with 4 articles, with 9 plumose setae on inner side and 2 setae on outer side, plus a simple small setae on apex. FIGURE 1. First zoea: A complete larvae lateral view; A’ carapace denticles; B antennule; C antenna; D mandibles; E maxillule; F maxilla; G first maxilliped; H second maxilliped; I third maxilliped; J telson. Scale bars: 0.1mm (A–D, F–J); 0.5mm (E). FIGURE 2. Second zoea: A complete larvae dorsal view; B antenna; C mandibles; D pleon lateral view; E first pereiopod; F fifth pereiopod. Third zoea: G antennule; H mandibles; I maxilla; J first pereiopod; K fifth pereiopod; L telson and uropods. Scale bars: 0.1mm. Mandible ( Figure 2C ): asymmetrical, both mandibles bearing 4 incisors and 1 lacinia mobilis , armature of molar processes as illustrated. Maxillule: unchanged beside size. Maxilla: unchanged beside size. First maxilliped: exopod entire with 1+1+2+1 plumose setae; otherwise unchanged besides size. Second maxilliped ( Figure 2A ): coxopodite unarmed; basipodite with 1+2+3 serrulate setae; exopod entire bearing 5 pairs of plumose setae on distal part; otherwise unchanged besides size. Third maxilliped ( Figure 2A ): coxopodite unarmed; basipodite with 2+1 serrulate setae; endopod with 4 articles with 2+1 simple, 0, 1+4 plumodenticulate, 2+1 (1 simple and 2 plumodenticulate) setae; exopod entire bearing 5 pairs of plumose setae on distal part. First pereiopod ( Figure 2E ): biramous bud. Second to fourth pereiopods: absent. Fifth pereiopod ( Figure 2F ): small bud. Pleon ( Figure 2A, 2D ): 1 pair of small setae on the dorso-posterior side of the 3 rd somite. Otherwise unchanged, besides size. Pleopods: absent. Uropods: absent. Telson ( Figure 2A ): triangular, indented medially with 8 + 8 posterior processes. Third Zoea ( Figure 2G–L ) Size: TL= 3.61± 0.10 mm ; CL= 0.86± 0.11 mm ; RL= 0.33± 0.05 mm ; N = 10 Carapace: rostrum reaching a quarter of antennular peduncle; otherwise unchanged. Antennule ( Figure 2G ): peduncle with 2 articles, with 2 small simple setae proximally near the stylocerite, 1 small plumose setae in a more distal position, and 2–3+1 distally; distal article bearing 2–3 plumose setae and 5–7 simple setae; endopod with 1 long plumose setae terminally; exopod with 1 plumose setae, 1–2 small plumose setae and 2–4 aesthetascs. Antenna: peduncle entire; endopod apically with 1 long plumose setae; exopod with 3 articles, with 11–12 plumose setae on inner side and 2 setae on outer side, plus a simple small setae on apex. Mandible ( Figure 2H ): asymmetrical, both mandibles bearing 4 incisors, 1 lacinia mobilis and 1 submarginal setae; armature of molar processes as illustrated. Maxillule: unchanged besides size. Maxilla ( Figure 2I ): coxopodite bilobed with 9–11 (4 serrulate and 5–7 pappose) and 4 (1 serrulate and 3 pappose) setae, respectively; basipodite bilobed, both bearing 4 (1 serrulate and 3 pappose) setae; endopod entire bearing 3 (2 serrulate and 1 simple) +2 serrulate +2 serrulate setae +1–2 serrulate setae; exopod with 8 marginal plumose setae bearing lateral microtricha. First maxilliped: unchanged besides size. Second maxilliped: unchanged besides size. Third maxilliped: coxopodite unarmed; basipodite with 2+1 serrulate setae; endopod with 4 articles with 2+2 simple, 1 simple, 1+4 plumodenticulate, 2+1 (1 simple and 2 plumodenticulate) setae. Otherwise unchanged besides size. First pereiopod ( Figure 2J ): functional, biramous; coxopodite unarmed; basipodite with 2 setae; endopod: first article with 1–2+1 simple setae, second article without setae, third article with 3–4 plumodenticulate setae, fourth article with 1+2 (1 simple and 2 plumodenticulate) setae; exopod entire with 4 pairs of plumose setae on distal part. Second to fourth pereiopods: absent Fifth pereiopod ( Figure 2K ): uniramous bud Pleon: 6 somites; 1 pair of small dorso-lateral spines on posterior margin of 5 th somite, 1 pair of small lateral spines on 6 th somite and 1 pair small posterior setae on 3 rd somite. Pleopods: absent Uropod ( Figure 2L ): biramous, smaller than telson; exopod developed with 11–12 plumose setae; endopod small with 2 small plumose setae apically. Telson ( Figure 2L ): first pair of processes in an anterolateral position, about a fifth of total length of telson; otherwise unchanged besides size. Fourth Zoea ( Figure 3A–D ) Size: TL= 3.87± 0.15 mm ; CL= 0.86± 0.08 mm ; RL= 0.33± 0.03 mm ; N = 10 Carapace ( Figure 3A ): rostrum reaching approximately a third of antennular peduncle; 1 supraorbital spine, 1 antenal spine, 1 pterigostomial spine and 4 denticles along ventral margin. Antennule ( Figure 3A ): peduncle with 2 articles, with 2 small simple setae proximally, 3 small plumose setae in a more distal position, and 4 plumose setae distally; distal article bearing 2–4 plumose setae and 3–5 simple setae; endopod with 1 long plumose setae; exopod with 1 plumose setae and 2 aesthetascs. Antenna ( Figure 3B ): peduncle entire; endopod apically with 1 long plumose setae; exopod entire, with 13–17 plumose setae on inner side and 1–2 setae on outer side, plus a simple small setae on apex. Mandible ( Figure 3C ): asymmetrical, both mandibles bearing 4 incisors, 1 lacinia mobilis and 2 submarginal setae; armature of molar processes as illustrated. Maxillule: unchanged besides size. Maxilla: exopod with 8–12 marginal plumose setae. Otherwise unchanged besides size. First maxilliped: unchanged besides size. Second maxilliped ( Figure 3A ): endopod with 3 articles with 3 (2 simple and 1 plumodenticulate) +1 simple, 1 simple + 1 plumodenticulate, 1+4+1 (3 simple and 3 plumodenticulate) setae; Otherwise unchanged besides size. Third maxilliped ( Figure 3A ): endopod with 4 articles with 2+2 simple, 1 simple, 1–3+4 plumodenticulate, 2+1 (1 simple and 2 plumodenticulate) setae; exopod entire bearing 6–7 pairs of plumose setae on distal part. Otherwise unchanged besides size. First pereiopod ( Figure 3A ): endopod: first article with 2+2 setae, second article with 1 plumodenticulate setae, third article with 1+4 plumodenticulate setae, fourth article with 2 simple and 1 plumodenticulate setae; exopod entire with 5–7 pairs of plumose setae on distal part; Otherwise unchanged besides size. Second pereiopod ( Figure 3A ): biramous bud. Third to fourth pereiopods: absent Fifth pereiopod ( Figure 3A, 3D ): developed, uniramous. Coxopodite unarmed; basipodite bears 1 simple setae; first article with 1 setae; second article with 0–1 setae; third article with 1 setae; fourth article flattened and paddlelike enlarged with margins serrated (6–7 and 5 teeth on each side respectively), bearing 2 setae distally; fifth article small with 2 terminal setae. Pleon ( Figure 3A ): 6 somites; 1 pair of small dorso-lateral spines on posterior margin of 5 th somite, 1 pair of small lateral spines on 6 th somite and 2 pairs small posterior setae on 3 rd somite; dorsal region of the 3 rd pleomere is slightly curved. Pleopods: absent Uropod ( Figure 3A ): biramous; exopod well developed reaching the end of telson, with 15–19 plumose setae bearing lateral microtricha; endopod with 7–12 plumose setae. Telson ( Figure 3A ): unchanged besides size. Fifth Zoea ( Figure 3E–I ) Size: TL= 4.30± 0.43 mm ; CL= 0.77± 0.38 mm ; RL= 0.30± 0.09 mm ; N = 10 Carapace: rostrum reaching a fifth of antennular peduncle; 2 denticles along ventral margin. Otherwise unchanged. Antennule ( Figure 3E ): peduncle with 2 articles; proximal article with 4–5 small simple setae on stylocerite and 1+1+2+3-4 plumose setae along the article; distal article with 4–6 plumose setae; endopod with 3 articles, distal article with 1 long plumose setae and 2–3 simple setae; exopod with 3 articles with 2 aesthetascs in the first one and 1 plumose +0–1 simple setae apically. Antenna: peduncle entire; endopod developing without setae; exopod entire, with 18 plumose setae on inner side, plus simple small setae on apex. Mandible: asymmetrical, both mandibles bearing 4 incisors and 1 lacinia mobilis , right mandible with 2 submarginal setae and left with 3 submarginal setae. Maxillule: coxopodite with 8 cuspidate setae, basipodite with 8 cuspidate setae bearing lateral microtricha; endopod with 2 strong subterminal serrulate setae and 3 strong terminal serrulate setae. Maxilla: exopod with 11–13 marginal plumose setae. Otherwise unchanged besides size. First maxilliped ( Figure 3F ): basipodite with 13 (4 serrulate and 9 pappose) setae; unchanged besides size. FIGURE 3. Fourth zoea: A complete larvae lateral view; B antenna; C mandibles; D detail of fifth’s pereiopod propodus and dactylus. Fifth zoea: E antennule; F first maxilliped; G second pereiopod; H third pereiopod; I detail of fifth’s pereiopod propodus and dactylus; J uropods and telson. Scale bars: 0.1mm. Second maxilliped: exopod entire bearing 4 pairs of plumose setae on distal part. Otherwise unchanged besides size. Third maxilliped: unchanged besides size. First pereiopod: endopod: first article with 2+3 setae, second article with 1 plumodenticulate setae, third article with 1+4 plumodenticulate setae, fourth article with 2 simple and 1 plumodenticulate setae; otherwise unchanged besides size. Second pereiopod ( Figure 3G ): biramous, functional; coxopodite unarmed; basipodite with 1 simple setae; endopod: first article with 2+3 setae, second article without setae, third article with 4 plumodenticulate, fourth article with 2 simple and 1 plumodenticulate setae; exopod entire with 4 pairs of plumose setae on distal part. Third pereiopod ( Figure 3H ): biramous bud. Fourth pereiopods: absent Fifth pereiopod ( Figure 3I ): coxopodite with 1 simple setae; basis with 1 simple setae; first article with 2 setae; second article with 2 setae and a spine-like process; third article margins serrated with teeth on both sides and bearing 3 setae; otherwise unchanged besides size. Pleon: 6 somites; 1 pair of small lateral spines on 6 th somite and 2 pairs small posterior setae on 3 rd somite. Pleopods: absent Uropod ( Figure 3J ): biramous, longer than telson; exopod with 22+1 plumose setae; endopod with 17 plumose setae. Telson ( Figure 3J ): margins laterally parallel; posterior margin concave with a central invagination. Sixth Zoea ( Figure 4A–E ) Size: TL= 4.18± 1.10 mm ; CL= 1.01± 0.63 mm ; RL= 0.33± 0.04 mm ; N = 3 Carapace: 1 denticle along antero-ventral margin; otherwise unchanged besides size. Antennule ( Figure 4A ): peduncle with 2 articles; proximal article with 4 small simple setae on stylocerite, 6 plumose setae along the inner margin, 1 simple setae and one row of plumose setae on outer posterior margin; distal article with a row of plumose setae on inner posterior margin; endopod with 6 articles, second and third articles with 1 aesthetasc each and, 1, 0, 2 setae in the distal articles, respectively; exopod with 4 articles with 3 aesthetascs in the second article and 1 plumose setae and 2 simple setae in the distal one. Antenna: exopod entire, with 21 plumose setae on inner side, 1 spine plus a simple small setae; otherwise unchanged besides size. Mandible ( Figure 4B ): asymmetrical, both mandibles bearing 4 incisors, 1 lacinia mobilis and 3 submarginal setae. Maxillule: coxopodite with 11 cuspidate setae. Otherwise unchanged besides size. Maxilla: coxopodite bilobed with 10 (4 serrulate and 6 pappose) and 4 (1 serrulate and 3 pappose) setae, respectively; basipodite bilobed, both bearing 4 (1 serrulate and 3 pappose) setae; endopod entire bearing 3 (2 serrulate and 1 simple) +2 serrulate +4 serrulate setae; exopod with 13 marginal plumose setae bearing lateral microtricha. First maxilliped: coxopodite with 2+1+2 serrulate setae; basipodite with 14 (4 serrulate and 10 pappose) setae; endopod with 4 articles with 3,1,2,3 pappose setae; exopod entire with 1+3 plumose setae. Second maxilliped: coxopodite unarmed; basipodite with 1+2+3 serrulate setae; exopod entire bearing 6 pairs of plumose setae on distal part. Otherwise unchanged. Third maxilliped: coxopodite unarmed; basipodite with 3 serrulate setae along inner margin; endopod with 4 articles with 2+2 simple, 1 simple, 2+4 plumodenticulate, 2+1 (1 simple and 2 plumodenticulate) setae; exopod entire bearing 8 pairs of plumose setae on distal part. First pereiopod: biramous; coxopodite with 1 simple setae; basipodite with 1 simple setae; endopod: first article with 2+2 setae, second article with 3 plumodenticulate setae, third article with 3+4 plumodenticulate setae, fourth article with 2 simple and 1 plumodenticulate setae; exopod entire with 8 pairs of plumose setae on distal part. Second pereiopod: biramous; coxopodite without setae; basipodite with 2 simple setae; endopod: first article with 2+1 setae, second article with 2 plumodenticulate, third article with 4 plumodenticulate, fourth article with 2 simple and 1 plumodenticulate setae; exopod entire with 6 pairs of plumose setae on distal part. Third pereiopod ( Figure 4C ): biramous; coxa without setae; basipodite with 1 simple setae; endopod: first article with 2 simple setae, second article without setae, third article without setae, fourth article with 5 plumodenticulate setae, fifth article with 2 plumodenticulate setae; exopod entire with 3 pairs of plumose setae on distal part. FIGURE 4. Sixth zoea: A antennule; B mandibles; C third pereiopod; D fourth pereiopod; E uropods and telson. Seventh zoea: F larvae dorsal view; G maxillule; H maxilla. Scale bars: 0.1mm. Fourth pereiopod ( Figure 4D ): biramous bud Fifth pereiopod: developed, uniramous. coxopodite without setae; basipodite bears 1 simple setae; endopod: first article without setae; second article with 1 setae and a spine-like process; third article with 1+2 setae and a spine-like process; fourth article paddle-like, margins 2/3 serrated with teeth on both sides and bearing 1+2+1 setae; fifth article small with 2 terminal setae. Pleon: unchanged besides size. Pleopods: absent Uropods ( Figure 4E ): biramous, longer than telson, with 2 base spines; exopod with 2 simple setae +1 spine +24 plumose setae +3 simple setae; endopod with 3 simple setae +18 plumose setae. Telson ( Figure 4E ): second pair of processes also in an anterolateral position; otherwise unchanged besides size. Seventh Zoea ( Figure 4F–H ) Size: TL= 5.46± 0.40 mm ; CL= 1.23± 0.15 mm ; RL= 0.56± 0.06 mm ; N = 10 Carapace ( Figure 4F ): rostrum with 1 spine in front of dorsal organ; anterio-ventral margin without denticles; otherwise unchanged besides size. Antennule ( Figure 4F ): peduncle with 2 articles; stylocerite with 5–7 small simple setae, proximal article with 3–4 setae on inner margin, 8–13 plumose setae along the inner margin of the article and 1 row of plumose setae on outer margin and posterior end; distal article with row of plumose setae terminally; endopod with 16–25 articles; exopod with 10–21 articles with 3 aesthetascs in the forth article and 1 plumose setae and 2 simple setae apically. Antenna ( Figure 4F ): endopod reaching about a third of exopod, with 3 small setae apically; exopod with 23–30 plumose setae on inner side, 1 spine plus 1–3 simple small setae on the outer margin. Mandible: both mandibles bearing 4 incisors, 1 lacinia mobilis and a molar process; right mandible with 2–6 submarginal setae and left with 3–5 submarginal setae. Maxillule ( Figure 4G ): coxopodite with 14–15 cuspidate setae, basipodite with 9 cuspidate setae bearing lateral microtricha; endopod with 2 strong subterminal serrulate setae and 3 strong terminal serrulate setae. Maxilla ( Figure 4H ): coxopodite bilobed with 14–15 (4 serrulate and 10 pappose) and 4 (1 serrulate and 3 pappose) setae, respectively; basipodite bilobed, bearing 6–7 (1 serrulate and 5 pappose) and 5–6 (1 serrulate and 4 pappose) setae; exopod with 19–30 marginal plumose setae bearing lateral microtricha; otherwise unchanged besides size. First maxilliped: basipodite with 17–18 (8 serrulate and 10 pappose) setae; endopod with 4 articles with 3,1– 2,2,4 pappose setae; exopod entire with 0–1 plumose setae on anterior outer margin and 1+4 plumose setae terminally; otherwise unchanged besides size. Second maxilliped ( Figure 4F ): endopod with 3 articles with 2–3 (1–2 simple +1 plumodenticulate) +1 simple, 2–3 (1–2 simple +1 plumodenticulate), 0–1+5 (2–3 simple +3 plumodenticulate) setae; otherwise unchanged. Third maxilliped ( Figure 4F ): endopod with 4 articles with 2+2 simple, 1 simple, 0–2 (0–1 simple +1 plumodenticulate) +2+4 plumodenticulate, 1–2 simple +2 plumodenticulate setae; otherwise unchanged besides size. First pereiopod ( Figure 4F ): coxopodite with 1–2 simple setae; basipodite with 0–3 simple setae; otherwise unchanged besides size. Second pereiopod ( Figure 4F ): basipodite with 2–3 simple setae; endopod: first article with 1–2+1–2 setae, second article with 2 plumodenticulate setae, third article with 4–5 plumodenticulate setae, fourth article with 2 simple and 1 plumodenticulate setae; exopod entire with 11–12 pairs of plumose setae on distal part; otherwise unchanged besides size. Third pereiopod ( Figure 4F ): basipodite with 2+0–1 simple setae; endopod: first article with 1–2 setae, second article with 1 simple setae and 1 spine-like process, third article with 1–3 setae, fourth article paddle-like bearing 11–18 sparsely plumodenticulate setae, fifth article small with 2 plumodenticulate and 1 simple terminal setae; exopod entire with 9–10 pairs of plumose setae on distal part; otherwise unchanged besides size. Fourth pereiopod ( Figure 4F ): biramous; coxopodite with 0–1 setae; basipodite with 1–2 simple setae; endopod: first article with 0–2 setae, second article with 1–2 setae, third article with 2–6 setae, fourth article enlarged, paddlelike, bearing 12–17 sparsely plumodenticulate setae, fifth article small with 2 simple and 1 simple terminal setae; exopod entire with 5–6 pairs of plumose setae. Fifth pereiopod ( Figure 4F ): coxopodite with 1 simple setae; basipodite bears 2 simple setae; first article with 0–2 setae; second article with 1–3 setae and a spine-like process; third article sparsely setose with 1–2 spine-like setae and a spine-like process; fourth article paddle-like, margins 2/3 serrated with teeth on both sides and bearing 5–15 sparsely plumodenticulate setae; fifth article small with 2 terminal setae. Pleon ( Figure 4F ): unchanged besides size. Pleopods: absent Uropods ( Figure 4F ): longer than telson; protopod with edged margins dorsally; exopod with 27–33 plumose setae along inner and posterior margin, 2–3 simple setae on anterior and 1 spine posteriorly on outer margin; endopod with 24–34 plumose setae. Telson ( Figure 4F ): margins slightly narrower posteriorly; otherwise unchanged besides size. Eighth Zoea ( Figure 5A–C ) Size: TL= 6.20± 2.57 mm ; CL= 1.67± 0.15 mm ; RL= 0.83± 0.09 mm ; N = 10 Carapace: unchanged besides size. Antennule ( Figure 5A ): stylocerite with 8 small simple setae, proximal article now with 3 setae on inner margin, 11–15 plumose setae along the margin of the segment, one row of plumose setae on outer margin and posterior end, 0–3 simple setae and 1 row of plumose setae on outer margin and posterior end, 2–4 simple setae 1 row of plumose setae terminally; endopod with 24–26 articles; exopod with 18–26 articles with 2–3 aesthetascs in the third article and 3–4 aesthetascs in the forth article and 1 plumose setae and 4–7 simple setae apically. Antenna ( Figure 5B ): peduncle with 2 simple setae; endopod with 2 articles with 1–2 simple setae on the margin of the 2 nd article and 5 small setae apically; exopod with 29–32 plumose setae on inner side, 1 spine plus 3–6 simple small setae. Mandible ( Figure 5C ): right mandible with 6–8 submarginal setae and left with 4–5 submarginal setae; otherwise unchanged besides size. Maxillule: coxopodite with 12–20 cuspidate setae, basipodite with 11–15 cuspidate setae bearing lateral microtricha; endopod with 2–3 strong subterminal serrulate setae and 3 strong terminal serrulate setae. Maxilla: coxopodite bilobed with 15 (4 serrulate and 10 pappose) and 4 (1 serrulate and 3 pappose) setae, respectively; basipodite bilobed, both bearing 8 (1 serrulate and 7 pappose) setae; endopod entire bearing 3 (2 serrulate and 1 simple) +2 serrulate +4 serrulate setae; exopod with 28–34 marginal plumose setae bearing lateral microtricha. First maxilliped: exopod with 1–4 plumose setae on anterior outer margin and 4–5 plumose setae terminally; otherwise unchanged besides size. Second maxilliped: unchanged besides size. Third maxilliped: unchanged besides size. First pereiopod: unchanged besides size. Second pereiopod: endopod: first article with 1–2+1–2 setae, second article with 1–4 setae and 2 plumodenticulate setae, third article with 4–5 plumodenticulate setae, fourth article with 2–3 simple and 1 plumodenticulate setae; exopod entire with 11–13 pairs of plumose setae on distal part; otherwise unchanged besides size. Third pereiopod: coxopodite with 1–2 simple setae; endopod: first article with 2–4 setae, second article with 0–2 setae and 1 spine-like process, third article with 2–4 setae, fourth article enlarged, paddle-like, bearing 16–22 sparsely plumodenticulate setae, fifth article small with 2 simple and 2–3 terminal plumodenticulate setae; exopod entire with 10–11 pairs of plumose setae on distal part. Otherwise unchanged besides size. Fourth pereiopod: coxopodite with 1 setae; basipodite with 1–2 simple setae; endopod: first article with 1–4 setae, second article with 1–3 setae, third article with 3–6 plumodenticulate setae, fourth article margins 1/3 serrated bearing 13–24 sparsely plumodenticulate setae, fifth article small with 2–4 simple and 1 terminal setae; exopod entire with 6 pairs of plumose setae on distal part. Otherwise unchanged besides size. Fifth pereiopod: developed, uniramous. Coxopodite with 0–1 simple setae; basipodite bears 1 simple setae; first article with 1–7 sparse setae; second article with 1–5 setae and 1 spine-like process; third article sparsely setose with 6 spine-like setae and 1 spine-like process; fourth article margins 2/3 serrated with teeth on both sides and bearing 14–31 sparsely plumodenticulate setae; fifth article small with 3–6 setae. Pleon: 6 somites; possibly with 3 small simple dorsal setae on the 4 th somite and 1–2 small dorsal setae on the 5 th somite. Pleopods: small buttons. FIGURE 5. Eighth zoea: A antennule; B antenna; C mandibles. Ninth zoea: D larvae dorsal view; E first maxilliped; F pleopods. Scale bars: 0.1mm. Uropods: exopod with 3–4 simple setae on anterior outer margin, 1–6 simple setae and 1 spine on posterior outer margin, followed by 32–40 plumose setae along posterior and inner margin; endopod with 33–38 plumose setae along margin. Otherwise unchanged besides size. Telson: slightly narrower posteriorly. Otherwise unchanged besides size. Ninth Zoea ( Figure 5D–F ) Size: TL= 7.58± 1.48 mm ; CL= 2.26±2.57 mm ; RL= 1.07 ± 0.17 mm ; N = 3 Carapace ( Figure 5D ): rostrum reaching about a third of antennular peduncle, one spine in front of the dorsal organ and 2 spines on the rostrum. Otherwise unchanged. Antennule ( Figure 5D ): stylocerite with 7–10 small simple setae, proximal article with 2–3 setae on inner margin, 16–20 plumose setae along the margin of the article +1 row of plumose setae +1–2 simple setae +1 row of plumose setae, 1–4 simple setae +1 row of plumose setae; endopod with 35–38 articles; exopod with 40–45 articles with 0–1 aesthetascs in the third article, 3–5 aesthetascs in the fourth, 4–6 aesthetascs in the fifth, 2–5 aesthetascs in the sixth, 0–3 aesthetascs in the seventh and 4–5 simple setae apically. Antenna ( Figure 5D ): endopod nearly twice the length of exopod, with 17–20 articles, each one with 0–3, except the terminal one with 3–5, simple setae; exopod entire, with 39–41 plumose setae on inner side, 1 spine plus 5–7 simple small setae. Mandible: right mandible with 9–10 submarginal setae and left with 7 submarginal setae. Otherwise unchanged besides size. Maxillule: coxopodite with 13–16 cuspidate setae, basipodite with 11–12 cuspidate setae; endopod with 3–4 strong subterminal serrulate setae and 3 strong terminal serrulate setae. Otherwise unchanged besides size. Maxilla: coxopodite bilobed with 18 (4 serrulate and 14 pappose) and 4 (1 serrulate and 3 pappose) setae, respectively; endopod entire bearing 3 (2 serrulate and 1 simple) + 2–3 serrulate +4 serrulate setae; exopod with 48 marginal plumose setae. Otherwise unchanged besides size. First maxilliped ( Figure 5E ): basipodite with 18–19 (8 serrulate and 10–11 pappose) setae; endopod with 4 articles with 2–3,1–3,2,3–4 pappose setae; exopod showing the caridean lobe with 9–10 plumose setae along the margin and with 6–8 pairs of plumose setae on distal part. Otherwise unchanged besides size. Second maxilliped ( Figure 5D ): basipodite with 6–8 serrulate setae along inner margin; endopod with 3 articles with 2–3+1 (2 simple + 1–2 plumodenticulate), 2–4 plumodenticulate, 1+5 (3 simple and 3 plumodenticulate) setae; exopod entire bearing 5–6 pairs of plumose setae on distal part. Otherwise unchanged besides size. Third maxilliped ( Figure 5D ): coxopodite with 0–1 simple small setae; endopod with 4 articles, with 2+1 simple, 1 simple and 2 plumodenticulate, 16–24 plumodenticulate, 3–4 (1–2 simple and 2 plumodenticulate) setae; exopod entire bearing 11–13 pairs of plumose setae on distal part. Otherwise unchanged besides size. First pereiopod ( Figure 5D ): endopod: first article with 2–3 setae, second article with 2 plumodenticulate setae, third article with 1–2 simple and 8 plumodenticulate setae, fourth article with 5 simple and 1 plumodenticulate setae; exopod entire with 18 pairs of plumose setae on distal part. Otherwise unchanged besides size. Second pereiopod ( Figure 5D ): coxopodite with 2–3 simple setae; endopod: first article with 4–5 setae, second article with 4 setae and 3 plumodenticulate setae, third article with 7–12 plumodenticulate setae, fourth article with 7 simple and 1 plumodenticulate setae; exopod entire with 23–25 pairs of plumose setae on distal part. Otherwise unchanged besides size. Third pereiopod ( Figure 5D ): coxopodite with 2–4 simple setae; endopod: first article with 5–9 setae, second article with 1–6 setae and 1 spine-like process, third article with 4–6 setae, fourth article bearing 26–33 sparsely plumodenticulate setae, fifth article small with 4–5 simple and 1 terminal setae; exopod entire with 14–15 pairs of plumose setae on distal part. Otherwise unchanged besides size. Fourth pereiopods ( Figure 5D ): coxopodite with 1–2 setae; basipodite with 3 simple setae along margin; endopod: first article with 5–6 setae, second article with 2–3 setae, third article with 10–15 plumodenticulate setae, fourth article bearing 34–40 sparsely plumodenticulate setae, fifth article small with 5 simple and 1 terminal setae; exopod entire with 10–11 pairs of plumose setae on distal part. Fifth pereiopod ( Figure 5D ): coxopodite with 1–3 simple setae; basipodite bears 1–3 simple setae; first article sparsely setose; second article with 4–7 spine-like setae and 1 spine-like process; third article sparsely setose with 11–16 spine-like setae and 1 spine-like process; fourth article bearing 21–27 sparsely plumodenticulate setae; Otherwise unchanged besides size. Pleon ( Figure 5D ): with 0–4 small simple setae along dorsal margin on the 1 st somite, 0–2 small simple setae along dorsal margin on the 3 rd somite, 2–5 small simple setae along dorsal margin on the 4 th somite, 1–2 small simple setae along dorsal margin and 0–1 simple setae on posterior ventral margin of the 5 th somite and 6 small simple setae along ventral margin the 6 th somite. Pleopods ( Figure 5D, 5F ): bilobed, endopod bud-like with 2–4 simple apical setae as figured; exopod as bud. Uropods ( Figure 5D ): exopod almost twice the length of telson with 6–10 simple setae on anterior outer margin, 6–8 simple setae and 1 spine on posterior outer margin, followed by 48–51 plumose setae along posterior and inner margin; endopod with 49–50 plumose setae along margin. Otherwise unchanged besides size. Telson ( Figure 5D ): unchanged besides size. Tenth Zoea ( Figure 6A–U ) Size: TL= 11.61± 1.62 mm ; CL= 3.17± 0.69 mm ; RL= 1.95 ± 0.49 mm ; N = 6 Carapace ( Figure 6A, 6B ): one spine in front of the dorsal organ, 3–4 spines on the rostrum dorsal part and 0–1 ventrally. Otherwise unchanged. Antennule ( Figure 6A, 6C ): peduncle with 2 articles; stylocerite with 12–17 small simple setae, proximal article with 4–6 setae on inner margin, 20–24 plumose setae along the margin of the article +1 row of plumose setae + 3 simple setae +1 row of plumose setae, 3–10 simple setae +1 row of plumose setae; endopod with 63–71 articles; exopod with 62–75 articles with 0–2 aesthetascs in the second article, 2–4 aesthetascs in the third article, 5–7 aesthetascs in the fourth, 6–7 aesthetascs in the fifth, 4–7 aesthetascs in the sixth, 3–7 aesthetascs in the seventh, 0–4 aesthetascs in the eighth, 0–3 aesthetascs in the ninth and 4–9 simple setae apically. Antenna ( Figure 6A, 6D ): basipodite with 4+4–7 simple setae; endopod three times the length of scaphocerite, with 49–59 articles, each one with 0–4, except the terminal one with 3–4 simple setae; exopod entire with 48–54 plumose setae on inner side, 1 spine plus 10–13 simple small setae. Mandible ( Figure 6E ): right mandible with 10–12 submarginal setae and left with 8–9 submarginal setae. Otherwise unchanged besides size. Maxillule ( Figure 6F ): coxopodite with 18–21 cuspidate setae, basipodite with 19–21 cuspidate setae bearing lateral microtricha; endopod with 4–5 strong subterminal serrulate setae and 3 strong terminal serrulate setae. Maxilla ( Figure 6G ): coxopodite with 20–21 (4 serrulate and 16–17 pappose) and 4 (1 serrulate and 3 pappose) setae, respectively; endopod entire bearing 3 (2 serrulate and 1 simple) +3–4+4 serrulate setae, and 0–1 small plumose setae on the ventral external side; exopod with 64–77 marginal plumose setae; microtricha as illustrated. Otherwise unchanged besides size. First maxilliped ( Figure 6H ): coxopodite with 4–7 serrulate setae; basipodite with 20–22 (8–11 serrulate and 11–14 pappose) setae and 5–11 simple setae scattered around; endopod with 4 articles with 3–5,4–6,2,4 pappose setae; exopod showing the caridean lobe with 15–18 plumose setae along the margin, entire, with 4 pairs of plumose setae on distal part; epipodite as bud. Second maxilliped ( Figure 6I ): coxopodite with 0–1 simple setae; basipodite with 8 serrulate setae along inner margin and 1–3 simple setae along outer margin; endopod with 3 articles with 2 simple +2 plumodenticulate +3–4 simple, 1 simple +6 plumodenticulate, 4–5 simple +3 plumodenticulate setae; exopod entire bearing 5–6 pairs of plumose setae on distal part. Third maxilliped ( Figure 6J ): coxopodite with 1–3 simple small setae; basipodite with 3–4 serrulate setae along inner margin; endopod with 4 articles with 3–6 simple, 1 simple and 3–5 plumodenticulate, 23–31 plumodenticulate, 5–6 (3–4 simple and 2 plumodenticulate) setae; exopod entire bearing 17–19 pairs of plumose setae on distal part. First pereiopod ( Figure 6K ): coxopodite with 0–3 simple setae; basipodite with 2–4 simple setae; endopod: first article with 7 setae, second article with 1–2 simple and 1+3 plumodenticulate setae, third article with 7 simple and 6 plumodenticulate setae and a forming chela about a 3 rd of fourth article’s size, fourth article with 10 simple and 1 plumodenticulate setae; exopod entire with 23–25 pairs of plumose setae on distal part. Second pereiopod ( Figure 6L ): basipodite with 1–4 simple setae; endopod: first article with 8–16 setae, second article with 8 setae and 6 plumodenticulate setae, third article with 7–9 plumodenticulate setae and a forming chela about a 3 rd and a half of the fourth size bearing 6–8 simple setae, fourth article with 7–11 simple and 1 plumodenticulate setae. Otherwise unchanged besides size. FIGURE 6. Tenth zoea: A complete larvae lateral view; B rostrum and carapace denticles; C antennule; D antenna; E mandibles; F maxillule; G maxilla; H first maxilliped; I second maxilliped; J third maxilliped; K first pereiopod; L second pereiopod; M third pereiopod; N fourth pereiopod; O fifth pereiopod; P first pleopod; Q second pleopod; R third pleopod; S fourth pleopod; T fifth pleopod; U telson and uropods. Scale bars: 0.1mm. Third pereiopod ( Figure 6M ): basipodite with 1–5 simple setae; endopod: first article with 7–9 setae, second article with 12–17 setae and 1 spine-like process, third article with 14–22 setae, fourth article bearing 57–74 sparsely plumodenticulate setae, fifth article small with 7–10 simple and 1 terminal setae; exopod entire with 20–23 pairs of plumose setae on distal part. Otherwise unchanged besides size. Fourth pereiopods ( Figure 6N ): coxopodite with 2–7 setae; basipodite with 1–3 setae along margin; endopod: first article sparsely setose with 8 setae, second article sparsely setose with 7 setae, third article sparsely setose with 18 plumodenticulate setae, fourth article margins 1/3 serrated bearing 20–61 sparsely plumodenticulate setae, fifth article small with 4–9 simple and 1 terminal setae; exopod entire with 13–16 pairs of plumose setae on distal part. Fifth pereiopod ( Figure 6O ): coxopodite with 2–3 simple setae; basipodite bears 2–4 simple setae; first article sparsely setose; second article sparsely setose with 4–7 spine-like setae and 1 spine-like process; third article sparsely setose with 3–6 spine-like setae and a spine-like process; fourth article margins 2/3 serrated with teeth on both sides and bearing 27–56 sparsely plumodenticulate setae; fifth article small with 5 simple terminal and 3 spinelike setae. Pleon: with 3–5 small simple setae along ventral margin on the 1 st somite, 2–6 small simple setae along ventral margin on the 3 rd somite, 1–4 small simple setae along ventral margin on the 4 th somite and 1–4 small simple setae along ventral margin on the 5 th somite. Pleopods ( Figure 6P, 6Q, 6R, 6S, 6T ): protopods without setae; endopods with 4–8 small spines and exopods with 8–16 small spines, as illustrated. Uropods ( Figure 6U ): exopod with 3–7 simple setae on anterior outer margin, 8–16 simple setae and 1 spine on posterior outer margin, followed by 60–72 plumose setae along posterior and inner margin; endopod with 2–4 simple setae on anterior outer margin, 5–7 simple setae and 58–69 plumose setae along margin with lateral microtricha. Otherwise unchanged besides size. Telson ( Figure 6U ): Unchanged besides size. Decapodite ( Figure 7A–U ) Size: TL= 26.19 mm ; CL= 7.23 mm ; RL= 4.38 mm ; N = 1 Carapace ( Figure 7A ): carapace smooth, rostrum straight reaching over a half of antennular peduncle, with one spine in front of the dorsal organ, 6 spines on the dorsal part and 0–3 ventrally. Antennule ( Figure 7A, 7 A’, 7B): basal article of the antennular peduncle with stylocerite reaching over 1/4 of this article; peduncle with 3 articles; lateral antennular flagellum with aesthetascs extending from the 3 rd article to 20 th article. Antenna ( Figure 7A, 7 A’, 7C): basipodite with 4 simple setae; endopod nine times the length of scaphocerite, with 147 articles, each one with 0–1, except the terminal one with 8 simple setae; exopod entire, with 67 plumose setae on inner side, 1 spine plus 2 simple small setae. Mandibles ( Figure 7D ): unequal, without palp; presenting incisor process, as figured. Maxillule ( Figure 7E ): coxopodite with 36 cuspidate setae, basipodite with 51 cuspidate setae; endopod with 2 strong terminal serrulate setae and 5 setae along outer margin. Maxilla ( Figure 7F ): coxopodite bearing 23 setae; basipodite bilobed, each bearing 25 setae and 29 setae respectively; endopod entire bearing 1 setae on inner side, 1 terminal setae and 1 setae along outer margin; scaphognathite well developed, very large and not very broad, with 115 plumose setae along the margin. First maxilliped ( Figure 7G ): coxopodite with 8 serrulate setae; basipodite with 62 setae; endopod with 4 articles with (0,2,0,3) pappose setae; exopod showing the caridean lobe, not distinctly separated with 38 plumose setae along the margin, entire, with 10 pairs of plumose setae on distal part; epipodite bilobed. Second maxilliped ( Figure 7H ): coxa with 7 simple setae and a large epipod laterally; basis with 10 setae along inner margin and 4 setae along outer margin; endopod stout, terminal article, distally elongated, with rows of long marginal and submarginal setae; ischium with 4 setae along inner margin and 6 simple setae along outer margin; merus with 3 setae; carpus short with 2 setae along the margin; propod with row of distributed setae; exopod entire bearing 13 pairs of plumose setae on distal part. Third maxilliped ( Figure 7I, 7 I’): with long and slender endopod, distal article with 6 spines distributed terminally; exopod short, slender, reaching more than 2/3 of the two proximal endopod articles, with 2 terminal setae and 1 setae near the base. FIGURE 7. Decapodite: A specimen lateral view; A’ complete specimen lateral view; B antennule; C antenna; D mandibles; E maxillule; F maxilla; G first maxilliped; H second maxilliped; I third maxilliped; J first pereiopod; J’ first pereiopod, detail of chelae; K second pereiopod; K’ second pereiopod, detail of chelae; L third pereiopod; L’ third pereiopod, detail of chelae; M fourth pereiopod; M’ fourth pereiopod, detail of chelae; N fifth pereiopod; N’ fifth pereiopod, detail of chelae; O first pleopod; P second pleopod; Q third pleopod; R fourth pleopod; S fifth pleopod; T telson; U uropods. Scale bars: 0.1mm. First pereiopod ( Figure 7J, 7 J’): all articles well differentiated and setose as figured; propodus with a simple chela, reaching beyond the end of scaphocerite; palm about 3.5 times as long as dactylus; carpus almost as long as merus; exopod entire with four setae on ventral posterior margin and four setae on distal part. Second pereiopod ( Figure 7K, 7 K’): carpus twice as long as merus, with indistinct articulation; merus with equal length as ischium, both entire; exopod entire with eight setae on ventral posterior margin and two setae on distal part. Third pereiopod ( Figure 7L, 7 L’): similar to fourth and fifth pereiopod, being the smallest in size; with unarmed ischium; merus with four stout ventrolateral spines, over twice as long as carpus; propodus slightly shorter than merus with setae along margins; dactylus about 1/10 length of propodus, biunguiculate, with three additional slender spines on flexor margin, terminal unguis longer than ventral unguis; exopod entire with four setae on distal part. Fourth pereiopod ( Figure 7M, 7 M’): with four stout spines along ventrolateral margin of merus, which is almost two times as long as carpus; propodus slightly longer than merus with setae along margins; dactylus similar to third pereiopod, as figured; exopod entire with nine setae on ventral posterior margin, one on distal part and five on dorsal posterior margin. Fifth pereiopod ( Figure 7N, 7 N’): merus with five stout spines along lateral margin; propodus slightly shorter than merus with spines and setae along margins; dactylus similar to third and fourth pereiopod, as figured. Pleopods ( Figure 7O, 7P, 7Q, 7R, 7S ): endopod of first pleopod about half as long as exopod with 20 setae. Other pleopods with endopod slightly smaller than exopod, both with plumose setae on margins. Appendix masculina on second pleopod about half the size of appendix interna , with four setae on posterior end. Uropods ( Figure 7U ): protopod with two lateral teeth with seven setae on margin; exopod sub equal to endopod length, both with plumose setae on margins. Telson ( Figure 7T ): about 1.3 times as long as sixth pleomere; lateral margins of telson with many setae on distal half and 4 setae (2 long ones and 2 short ones) in the middle at proximal; dorsal surface with two pairs of spines, the first pair on 0.4 and second one on 0.8 of telson; posterior margins with a pair of short acute spines, two pairs of long acute ones, a pair of strong spines, a pair of short strong spines and a pair of long acute ones.