taxonID	type	description	language	source
03FA8E1EAB44FF8F471FF8E1FD69F8DF.taxon	materials_examined	Type species: Petrarca bathyactidis Fowler, 1889.	en	Kolbasov, Gregory A., Savchenko, Alexandra S., Yu, Meng-Chen, Tsao, Yao-Feng, Ganmanee, Monthon, Chan, Benny K. K. (2023): Integrative taxonomy, larval biology and functional morphology of the little known gall-forming coral endoparasite Petrarca (Thecostraca: Ascothoracida). Zoological Journal of the Linnean Society 198: 767-801, DOI: 10.1093/zoolinnean/zlad009
03FA8E1EAB44FF8F4581FF0AFB73FEB0.taxon	description	(FIGS 1 F, 2 – 7)	en	Kolbasov, Gregory A., Savchenko, Alexandra S., Yu, Meng-Chen, Tsao, Yao-Feng, Ganmanee, Monthon, Chan, Benny K. K. (2023): Integrative taxonomy, larval biology and functional morphology of the little known gall-forming coral endoparasite Petrarca (Thecostraca: Ascothoracida). Zoological Journal of the Linnean Society 198: 767-801, DOI: 10.1093/zoolinnean/zlad009
03FA8E1EAB44FF81453EFEDBFDA3FE20.taxon	materials_examined	Material examined: Four specimens in Astreopora sp., two specimens in Turbinaria mesenterina (Lamarck, 1816) and 12 specimens in Turbinaria sp., 22 ° 40 ʹ 39.2 ″ N, 121 ° 28 ʹ 57.2 ″ E, Green Island (L ǜ d ǎ o), Taiwan, 24.04.2015, at 4 – 20 m depth. Fifteen specimens in Turbinaria frondens (Dana, 1846) and four specimens in Turbinaria sp., 10 ° 41 ʹ 56.5 ˝ N, 99 ° 24 ʹ 28.4 ″ E, Koh Khai Island, Chumphon, Thailand, 06 – 07.07.2019, at 3 – 4 m depth. Grygier (1991 a) provided a detailed description of this species. We provide additional observations obtained mainly with SEM. Diagnosis: Carapace valves of adults with crenulated margins formed by protruding outer ends of five to eight radially directed dorsal and posterior irregular ridges; ridge exterior with irregular lumpy inflations; ventral side of carapace with few papillae; lateral surface of carapace without small papillae. Labrum with short lateral extensions; maxillae mostly exposed; mandibles with 15 – 25 sharp, simple or bifid teeth; teeth of the maxillules in lower half with spiniform tips and tending to form two rows; six pairs of thoracopods, first thoracopod setiform; penis with relatively small, squarish rami. Description: Living specimens pink or crimson-coloured (Fig. 1 F); tip of penis often extending out of carapace (Fig. 2 C, E, F). Adult (mature) specimens 5.26 – 6.08 mm long, 4.32 – 6.15 mm high (Figs 2 A – D, 5 A); young (juvenile) specimens (Fig. 2 E – G) 2.80 – 3.83 mm long and 2.40 – 3.33 mm high. Carapace (Figs 2, 5 A) roughly ovoid (spherical), valves with short anteriodorsal hinge line, lateral surfaces with five to eight radial ridges with irregular lumpy inflations. These inflations are less developed in the anterior part of adults and in young specimens. Dorsal and posterior margins of valve crenulated due to carapace ridges (Figs 2, 5 F). Ventral margin with few inconspicuous papillae (Fig. 5 C, D) with central micropore on tip (Fig. 5 E). Cuticle on lateral, external and internal surfaces of carapace with dense, polygonal, small swellings or bumps but without papillae, teeth or ctenes (Fig. 5 B, G). Body inflated, crescent-like, enclosed between carapace valves (Fig. 5 F). Cephalon with large adductor muscle lying above big oral cone (Fig. 5 F) flanked by five-segmented antennules. Thorax with arched dorsal margin but without distinct segmentation, with clusters of rudimentary uniramous thoracopods; cuticle of thorax with ctenoid scales in dorsal part (Fig. 5 H). Abdomen with massive first segment bearing long penis and vestigial rear part (Figs 5 F, 7 D). Antennules somewhat prehensile, with little armament or external sculpture on two distal segments (Figs 3 A – D, 6 A – D). First segment irregularly rectangular, narrowing somewhat distally; second segment trapezoidal; third segment almost triangular, narrowing toward lower / ventral margin; fourth segment slightly longer than wide, with slightly curved ventral margin, short distal seta inserted at anteriodorsal corner (Figs 3 A, D, 6 A, B). Fifth segment rectangular, slightly shorter and narrower than fourth and armed with sensory and grasping structures (Figs 3 A – C, 6 A, C, D). Short but massive curved claw with smooth concave margin arising from distal end of segment. Three rudimentary setae at base of claw and to each side; tiny pores (three to four) on inner and outer lateral sides. Claw sheathed by large, oval claw guard on posteriodistal corner; claw guard with three vestigial, distal setae with a terminal pore; developed aesthetasc almost half as long as claw guard, inserted closer to its distal end and terminating with four outgrowths, rudimentary seta at base of aesthetasc (Fig. 6 A, D). Ventral (postaxial) margin of fifth segment and claw guard with wrinkled cuticle bearing tiny pores (Fig. 6 C). Oral cone prominent, consisting of massive labrum underlaid posteriorly with massive, fused maxillae, unpaired medial languette and paired mandibles and maxillules (Figs 3 E – J, 5 F, 6 E – H). Labrum prow-shaped, with short posteriolateral extensions leaving maxillae largely exposed, exterior with dense ctenoid scales (Figs 3 E, 6 E). Medial languette with groups of sharp denticles (two to five) along anterior margin and dense basal pores forming ‘ pore-field’ (Fig. 6 F). Mandibles (Figs 3 F, G, 6 G) elongated, outer / upper margin with hillock and clusters of dense, thin setae in middle part, cutting edge straight, with 15 – 21 sharp, simple or rarely bifid teeth and one to two small spinules in middle part. Maxillules (Figs 3 H, I, 6 H) with sclerotized, triangular distal parts, inner margin with 17 – 20 teeth, teeth in upper half with often blunt tips, while those in lower half smaller, irregular, with spiniform tips and tending to form two irregular rows. Fused maxillae (Fig. 3 J) ending in a pair of short, rounded lobes, with dense ctenoid scales on lateral surfaces (Fig. 7 B), distal ends covered with sclerotized, thick and wrinkled cuticle without denticles, pores or setules (Fig. 7 A). Thoracopods 2 – 6 (Figs 4 B, 5 F) are present in all examined specimens, uniramous, unsegmented, grouped in cluster and arranged in an unorderly sequence in lateral view. Thoracopods 2 – 4 are conical with wide bases, while thoracopods 5 and 6 are shorter and narrower. Cuticle of thoracopods 2 – 6 has dense and long ctenoid scales (Fig. 7 C). Batteries of numerous ampuliform seminal receptacles (Fig. 4 B) associated with thoracopods 2 – 5 (~ 7, 20, 20 and 11 receptacles, respectively). Setiform thoracopod 1 (Fig. 4 A) present in all studied specimens. Long, thick, terminally bifid penis originates from large first abdominal segment (Figs 4 D, 5 F, 7 E). Long shaft of penis supports two small, squarish rami about 190 – 200 µm long (Figs 4 D, 7 E). Distal half of penis, including rami, bearing numerous short but wide conical setae. Rest of abdomen vestigial, apparently consisting of two segments (Figs 4 C, 5 F, 7 D). Posterior end of abdomen with cleavage that may represent rudimentary furcal rami (Fig. 7 D). Remarks: The morphology of our specimens generally corresponds to that of the individuals described by Grygier (1991 a), but differs in having rare ventral papillae on the carapace (absent in Grygier’s specimens) and presence of all six pairs of thoracopods (inconsistent number of thoracopods in Grygier’s material). However, these details could have been overlooked in the type material. This is the first finding of Petrarca in scleractinian corals of the genus Astreopora.	en	Kolbasov, Gregory A., Savchenko, Alexandra S., Yu, Meng-Chen, Tsao, Yao-Feng, Ganmanee, Monthon, Chan, Benny K. K. (2023): Integrative taxonomy, larval biology and functional morphology of the little known gall-forming coral endoparasite Petrarca (Thecostraca: Ascothoracida). Zoological Journal of the Linnean Society 198: 767-801, DOI: 10.1093/zoolinnean/zlad009
03FA8E1EAB4AFF8646E9FE79FDDDF973.taxon	description	(FIGS 1 G, 8 – 13)	en	Kolbasov, Gregory A., Savchenko, Alexandra S., Yu, Meng-Chen, Tsao, Yao-Feng, Ganmanee, Monthon, Chan, Benny K. K. (2023): Integrative taxonomy, larval biology and functional morphology of the little known gall-forming coral endoparasite Petrarca (Thecostraca: Ascothoracida). Zoological Journal of the Linnean Society 198: 767-801, DOI: 10.1093/zoolinnean/zlad009
03FA8E1EAB4AFF8646E9FE79FDDDF973.taxon	materials_examined	Material examined: Four specimens in Turbinaria bifrons Brüggemann, 1877, 22 ° 40 ʹ 39.2 ″ N, 121 ° 28 ʹ 57.2 ″ E, Green Island (L ǜ d ǎ o), Taiwan, 24.04.2015, at a depth of 4 – 20 m. Here we provide additional descriptions based on SEM analysis that supplements Grygier’s (1985) original description. Diagnosis: Carapace spherical, with large lumpy inflations, without radial ridges; carapace margin not crenulated; ventral side of carapace with large conical papillae, lateral surface with small papillae. Mandibles with 10 – 14 sharp, simple teeth; maxillules with small denticles; six pairs of thoracopods, first thoracopod setiform; penis with relatively small, rounded rami. Description: Living specimens pink, with dark inclusions at the anterior part of carapace (Fig. 1 G); tip of penis often extending out of carapace (Fig. 8). Adult (mature) specimens 2.13 – 2.42 mm long (1.2 – 3.8 mm from the Grygier material), 1.85 – 1.96 mm high (0.9 – 4.0 mm from the Grygier material), 2.00 – 2.24 mm wide (1.0 – 4.6 mm from the Grygier material) (Figs 8, 11 A). Carapace (Figs 8, 11 A) spherical; valves with conspicuous rounded lumpy inflations (8 – 14), without radial ridges, dorsal and posterior margins not crenulated; long, conical papillae with a tiny terminal pore on anterior half of ventral margin (Fig. 11 A, D). Cuticle on the lateral external and posteriodorsal internal surfaces of carapace with dense, polygonal, small swellings or bumps (Fig. 11 B, H). Small volcano-shaped papillae with tiny a terminal pore and microscopic pores inserted between bumps on external surface of carapace (Fig. 11 B, C, E). Thin cuticular lining (mantle) of inner surface of carapace with rows of dense ctenoid scales in central part (Fig. 11 F, G, I). Body inflated, curved, enclosed between carapace valves, tip of penis almost adjoins oral cone (Figs 8, 12 A). Cephalon with large adductor muscle lying above big oral cone flanked by five-segmented antennules. Thorax with sinusoid arched dorsal margin, without distinct segmentation, with clusters of rudimentary uniramous thoracopods (Fig. 12 A). Abdomen with massive first segment bearing long penis and vestigial rear part (Fig. 12 A). scales on lateral surface enlarged in circle outline). F, medial languette, lateral view (groups of denticles on anterior margin enlarged in oval outline, basal ‘ pore-field’ enlarged in circle outline). G, distal part of mandible (lower part of cutting edge enlarged in oval outline). H, maxillule (lower half of cutting edge enlarged in oval outline). Abbreviations: ae, aesthetasc; cl, claw; clg, claw guard. Scale bars in µm. Antennules W-shaped, with little armament (external sculpture) on two distal segments (Figs 9 A – C, 12 B – H). First segment irregularly rectangular, narrowing somewhat distally; second segment rectangular; third segment triangular, narrowing toward lower / ventral margin; fourth segment trapezoid, with slightly curved ventral margin, short distal seta inserted at anteriodorsal corner (Fig. 12 C). Fifth segment narrow, rectangular, slightly shorter than fourth and armed with sensory and grasping structures (Figs 9 B, C, 12 D – H). Short massive curved claw with smooth concave margin. Three rudimentary setae at base of claw and to each side (Figs 9 B, C, 12 F); tiny pores (five to seven) on inner and outer lateral sides (Figs 9 B, C, 12 D, E). Claw sheathed by large, oval claw guard (Fig. 12 F); claw guard with three vestigial, distal setae with a terminal pore (Fig. 12 H); developed subdistal aesthetasc almost half as long as claw guard, terminates with two outgrowths, rudimentary seta with a terminal pore at base of aesthetasc (Fig. 12 F, H). Ventral (postaxial) margin of fifth segment and claw guard with wrinkled cuticle bearing tiny pores (Fig. 12 G). Oral cone prominent (Figs 9 D, 12 A, I); labrum prow-shaped, exterior with dense ctenoid scales (Fig. 12 I, J); mandibles elongated, cutting edge straight, with 14 sharp, simple teeth (ten in Grygier specimen) (Fig. 9 E); maxillules with sclerotized, triangular distal parts, inner margin with small denticles (Fig. 9 F, G); fused maxillae ending in a pair of short, rounded lobes, with dense ctenoid scales on lateral surfaces (Figs 9 H, 12 I). Studied specimens with six pairs of thoracopods; thoracopod 1 setiform (Fig. 10 A); thoracopods 2 – 6 (Fig. 10 B) uniramous, unsegmented, grouped in cluster and arranged in an unorderly sequence in lateral view; thoracopods 2 – 4 longer and wider than thoracopods 5 and 6. Cuticle of thoracopods 2 – 6 with rows of dense and long ctenoid scales (Fig. 13 A, B). Batteries of ampuliform seminal receptacles (Fig. 10 B) associated with thoracopods 2 – 5 (~ 11, 6, 8 and 7 receptacles, respectively). Long, terminally bifid penis originates from large first abdominal segment (Figs 10 D, 12 A, 13 C). Shaft of penis supports two small, rounded rami about 125 µm long; distal part of penis, including rami, bearing numerous short but wide conical setae. Rest of abdomen vestigial, consisting of two segments bearing rare ctenoid scales and tiny pores; posterior end of abdomen with cleavage with few terminal denticles (Figs 10 C, 13 D – F). Remarks: Grygier & Nojima (1995) indicated that both P. morula and P. goanna may represent ‘ the extremes of morphological variability in a single species’. However, along with molecular evidence provided here, there are a number of morphological features that can be used to distinguish these species. Thus, P. morula differs from P. goanna in having numerous long conical papillae on the anterioventral part of carapace vs. only a few inconspicuous ventral papillae in P. goanna; exterior of carapace with small volcano-shaped papillae (absent in P. goanna) but without radial ribs and crenulated margin (present in P. goanna); teeth of the mandible are simple and fewer (simple and bifid teeth present in P. goanna); inner margin of maxillules with small denticles (bigger teeth in P. goanna); subterminal aesthetasc of claw guard terminates with two outgrowths (four in P. goanna).	en	Kolbasov, Gregory A., Savchenko, Alexandra S., Yu, Meng-Chen, Tsao, Yao-Feng, Ganmanee, Monthon, Chan, Benny K. K. (2023): Integrative taxonomy, larval biology and functional morphology of the little known gall-forming coral endoparasite Petrarca (Thecostraca: Ascothoracida). Zoological Journal of the Linnean Society 198: 767-801, DOI: 10.1093/zoolinnean/zlad009
03FA8E1EAB4DFF98473CF927FA7AFBBA.taxon	description	(FIGS 1 H, 14 – 18) Probably as P. morula in Grygier & Nojima, 1995: 93 – 96, fig. 18. Z o o b a n k R e g i s t r a t i o n: u r n: l s i d: z o o b a n k. o r g: a c t: A E C 7 C 3 E F - C 4 0 5 - 4 4 B 0 - B 0 6 9 - 364 D 18 F 6 B 6 A 3 Material examined: Two specimens (including holotype) in Turbinaria mesenterina, 22 ° 40 ′ 39.2 ″ N, 121 ° 28 ′ 57.2 ″ E, Green Island (L ǜ d ǎ o), Taiwan, 24.04.2015, at 4 – 20 m depth. Six specimens in Turbinaria sp., 25 ° 7 ʹ 49 ″ N, 121 ° 51 ʹ 26 ″ E, Keelung, Taiwan, 17.08.2015, at 4 – 10 m depth. Glycerin slide of the holotype (no. Mg 1248) with the carapace valves, the dissected mouth parts, antennules and rest of the body proper with thoracopods and penis, and SEM stub of paratype (no. Mg 1250) with the carapace valves and the body proper are deposited in the Zoological Museum of Moscow State University in Moscow, Russian Federation. Diagnosis: Carapace roughly ovoid, valves with eight to ten coalesced lumpy inflations, without distinct radial ridges; carapace margin uneven but not crenulated; ventral side of carapace with numerous, large conical papillae, lateral surface with small papillae. Mandible with 15 sharp, simple teeth; maxillules with 13 – 15 teeth; six pairs of thoracopods, first thoracopod setiform; penis with small, rounded rami. Etymology: The species is named after Dr Yoko Nozawa, our colleague and friend, in appreciation of his help with identification of coral hosts for our Ascothoracida studies. Description: Living specimens pink (Fig. 1 H). Adult (mature) specimens 3.70 – 4.22 mm long, 3.52 – 3.60 mm high and 3.2 – 3.3 mm wide (Figs 14, 17 A, G). Carapace (Fig. 14, 17 A, G) roughly ovoid, narrowing towards anterior end; each valve with eight to ten coalesced irregular lumpy inflations but without distinct radial ridges; dorsal and posterior margins of carapace uneven but not crenulated; anterioventral margins slightly curved or straight, with numerous, large conical papillae (Figs 14 C, E, 17 A, D); lateral surfaces and posterioventral margins with small papillae (Fig. 17 B, E). Carapace papillae volcano-shaped, terminating at central microscopic pore (Fig. 17 C, F). Cuticle of carapace with dense, polygonal, small swellings or bumps (Fig. 17 B, E, D). Body inflated, curved, enclosed between carapace valves, tip of penis close to oral cone (Figs 14 F, 17 H). Cephalon with large adductor muscle lying above big oral cone flanked by five-segmented antennules. Thorax with sinusoid arched dorsal margin, with hump in posterior part but without distinct segmentation, with cluster of rudimentary uniramous thoracopods (Figs 14 F, 17 H). Abdomen with massive first segment bearing long penis and vestigial rear part (Figs 14 E, F, 17 H). Antennules somewhat W-shaped and prehensile, with little armament of external sculpture on two distal cuticle with tiny pores on postaxial (ventral) margin of fifth antennular segment. H, distal parts of claw and claw guard, setae of claw guard and aesthetasc indicated by arrowheads. I, oral cone, lateroventral view. J, ctenoid scales on lateral surface of labrum. Abbreviations: ae, aesthetasc; cl, claw; clg, claw guard; lb, labrum; mo, mouth opening; mx 2, maxillae; oc, oral cone; pe, penis; rab, rudimentary abdomen; thp, thoracopods. Scale bars in µm. segments (Figs 14 F, 15 A – D). First segment irregularly rectangular, narrowing somewhat distally; second segment trapezoidal; third segment almost triangular, narrowing toward lower / ventral margin; fourth segment longer than wide, with curved ventral margin, short distal seta inserted at anteriodorsal corner (Fig. 15 A, B). Fifth segment rectangular, narrowing towards distal end, shorter and narrower than fourth and armed with sensory and grasping structures, ventral / postaxial margin almost straight, dorsal / preaxial margin concave (Fig. 15 C, D). Short, massive curved claw with smooth concave margin arising from distal end of segment. Three rudimentary setae at base of claw and to each side (Fig. 15 C, D); tiny pores (three to five) on inner and outer lateral sides (Fig. 15 C, D). Claw sheathed by large, hood-shaped, oval claw guard on posteriodistal corner; claw guard with three vestigial, distal setae; developed subterminal aesthetasc half as long as claw guard, terminates with two outgrowths (Fig. 15 C, D). Oral cone prominent (Figs 17 H, 18 A). Massive prow-shaped labrum with short posteriolateral extensions, leaving maxillae largely exposed, dense ctenoid scales on the exterior (Fig. 18 A, B). Mandibles (Fig. 15 E) elongated, cutting edge straight, with 15 sharp, simple teeth. Maxillules (Fig. 15 F, G) with sclerotized, triangular distal parts, inner margin with 13 – 16 teeth, teeth in upper half with blunt tips, while those in lower half smaller and spiniform. Fused maxillae (Figs 15 H, 18 A, B) with dense ctenoid scales on lateral surfaces, distal ends with rounded zones of sclerotized, thick and wrinkled cuticle without denticles, pores or setules. Six pairs of thoracopods; thoracopods 2 – 6 uniramous, unsegmented, grouped in cluster and arranged in an unorderly sequence in lateral view (Figs 16 B, 17 H). Thoracopod 1 setiform (Fig. 16 A); thoracopods 2 – 4 conical, longer and wider than thoracopods 5 and 6. Batteries of ampuliform seminal receptacles (Fig. 16 B) associated with thoracopods 2 – 5 (~ 14, 11 – 12, 9 – 10 and 9 receptacles, respectively). Cuticle of thoracopods 2 – 6 with dense and long ctenoid scales (Fig. 18 D). Long (~ 1.2 mm) and massive, terminally bifid penis originates from large first abdominal segment (Fig. 17 H). Shaft of penis supports two small, flat and rounded rami about 100 µm long; distal part of penis, including rami, bearing numerous short but wide conical setae (Figs 16 D, 18 E). Rest of abdomen vestigial, consisting of two indistinct segments; posterior end of abdomen with shallow cleft (Figs 16 C, 18 C). Remarks: The new species Petrarca nozaaeai sp. nov. is similar to the other two congeners, P. morula and P. goanna in utilizing Turbinaria corals as hosts. These species are characterized by the large lumpy inflations of the carapace, which are absent in other species of Petrarca. Petrarca nozaaeai differs from P. morula in having ovoid but not a spherical carapace and larger teeth on the inner margin of maxillules (small denticles in P. morula). Petrarca nozaaeai is distinguished from P. goanna: (1) by the absence of radial ribs and the crenulated margin of the carapace; (2) by numerous large, long conical papillae on the anterioventral part of the carapace; and (3) by simple and fewer teeth on the mandible (simple and bifid teeth present in P. goanna).	en	Kolbasov, Gregory A., Savchenko, Alexandra S., Yu, Meng-Chen, Tsao, Yao-Feng, Ganmanee, Monthon, Chan, Benny K. K. (2023): Integrative taxonomy, larval biology and functional morphology of the little known gall-forming coral endoparasite Petrarca (Thecostraca: Ascothoracida). Zoological Journal of the Linnean Society 198: 767-801, DOI: 10.1093/zoolinnean/zlad009
03FA8E1EAB53FF9345C7FB89FB29F9FA.taxon	description	(FIGS 1 I, 19 – 25) Z o o b a n k R e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org: act: BA 716 B 70 - D 5 B 8 - 49 A 1 - AF 02 - 670876 B 5 E 887 Material examined: Four specimens (including holotype) in Turbinaria bifrons and Turbinaria sp., 22 ° 40 ′ 39.2 ″ N, 121 ° 28 ′ 57.2 ″ E, Green Island (L ǜ d ǎ o), Taiwan, 24.04.2015, at 4 – 20 m depth. Glycerin slide of the dissected holotype (no. Mg 1249) with the carapace valves, antennules, mouth parts, and rest of the body with thoracopods and penis, and SEM stub of paratype (no. Mg 1251) with the carapace valves and the body are deposited in the Zoological Museum of Moscow State University in Moscow, Russian Federation. Diagnosis: Carapace roughly ovoid, carapace valves of adults with crenulated margins formed by protruding outer ends of six to seven radially directed dorsal and posterior irregular ridges; ridge exterior with irregular lumpy inflations; ventral side of carapace with numerous, large conical papillae, lateral surface with small papillae. Mandibles with 20 sharp, simple and bifid teeth; teeth of the maxillules in lower half with spiniform tips and tending to form two rows; six pairs of thoracopods, first thoracopod setiform; penis with relatively small, rounded cylindrical rami. Etymology. From Latin rubus, raspberry (Rubus idaeus L.), referring to the shape and colour of t h e c a r a p a c e i n l i v i n g i n d i v i d u a l s o f t h e n e w species resembling ripe raspberries. It is a noun in apposition. Description: Living specimens pink or crimson-coloured (Fig. 1 I). Adult (mature) specimens 4.35 – 5.71 mm long, 4.43 – 5.65 mm high (Figs 19 A – C, 22 A); young (juvenile) specimen (Fig. 19 D) 1.52 mm long and 1.54 mm high. Carapace roughly ovoid in adults (Figs 19 A – C, 22 A) and spherical in young specimen (Fig. 19 D); valves with short anteriodorsal hinge line; lateral surfaces with six to seven radial ridges with irregular lumpy inflations. These inflations are less developed in the anterior part of adults and in young specimens. Dorsal and posterior margins of valve crenulated due to carapace ridges (Figs 19 A – C, 22 A, E). Ventral margin curved, with numerous, large conical papillae (Fig. 22 C) with a central micropore at the tip (Fig. 22 D). Cuticle on the lateral external and internal surfaces of carapace with dense, polygonal, small swellings or bumps, with papillae on the external side and tiny pores on the internal side (Fig. 22 B, F). Body inflated, curved, enclosed between carapace valves, tip of penis close to oral cone (Fig. 22 E). Cephalon with large adductor muscle lying above big oral cone flanked by five-segmented antennules. Thorax with arched dorsal margin, with small hump in posterior part but without distinct segmentation, with clusters of rudimentary uniramous thoracopods (Fig. 22 E). Abdomen with massive first segment bearing long penis and vestigial rear part (Fig. 22 E). Antennules somewhat prehensile, with little armament (external sculpture) on two distal segments (Figs 20 A, 23 A). First segment irregularly rectangular, narrowing somewhat distally; second segment rectangular; third segment almost triangular, narrowing toward lower / ventral margin; fourth segment slightly longer than wide, with slightly curved ventral margin, short distal seta inserted at anteriodorsal corner (Figs 20 A, D, 23 A, B). Fifth segment rectangular, narrowing towards distal end, shorter and narrower than fourth and armed with sensory and grasping structures, ventral / postaxial margin uneven, dorsal / preaxial margin concave (Figs 20 B, C, 23 A, C – I). Short, massive curved claw with smooth concave margin arising from distal end of segment. Three rudimentary setae at base of claw and to each side (Figs 20 B, C, 23 E); tiny pores (six to eight) on inner and outer lateral sides (Figs 20 B, C, 23 C, D). Claw sheathed by large, hood-shaped, oval claw guard on posteriodistal corner (Figs 20 B, C, 23 A); claw guard with three vestigial, distal setae each with a terminal pore (Fig. 23 H); developed subterminal aesthetasc almost half as long as claw guard, terminating with three long outgrowths and rudimentary subterminal seta; rudimentary seta at base of aesthetasc (Fig. 23 F, G). Ventral (postaxial) margin of fifth segment and claw guard with wrinkled cuticle bearing tiny pores (Fig. 23 I). Oral cone prominent, consisting of massive labrum underlaid posteriorly with massive, fused maxillae, unpaired medial languette and paired mandibles and maxillules (Figs 20 E – I, 22 E, 24). Massive prow-shaped labrum with short posteriolateral extensions, leaving maxillae largely exposed, dense ctenoid scales on the exterior (Fig. 24 A). Mandibles (Figs 20 F, G, 24 B – D) elongated, outer / upper margin with dense, thin setae in the middle, cutting edge straight, with 20 sharp, simple or rarely bifid teeth and three small spinules (or reduced setae?) in the middle. Maxillules (Figs 20 H, 24 E – H) with sclerotized, triangular distal parts, inner margin with ~ 27 teeth, teeth in upper half with blunt tips, while those in lower half smaller, irregular, with spiniform tips and tending to form two irregular rows; dense basal pores forming ‘ pore-field’ (Fig. 24 H). Fused maxillae (Figs 20 I, 24 I – K) with dense ctenoid scales on lateral surfaces (Fig. 24 K), distal ends with rounded zones of sclerotized, thick and wrinkled cuticle without denticles, pores or setules (Fig. 24 J). Six pairs of thoracopods; thoracopods 2 – 6 uniramous, unsegmented, grouped in cluster and arranged in an unorderly sequence in lateral view (Figs 21 B, 22 E). Thoracopod 1 setiform (Figs 21 A, 25 A); thoracopods 2 – 4 conical, longer and wider than thoracopods 5 and 6. Batteries of ampuliform seminal receptacles (Fig. 21 B) associated with thoracopods 2 – 5 (~ 18 – 20, 20 – 22, 12 and 9 receptacles, respectively). Cuticle of thoracopods 2 – 6 with dense and long ctenoid scales (Fig. 25 B). Long (~ 0.8 – 1.0 mm) and massive, terminally bifid penis originates from large first abdominal segment (Fig. 22 E). Shaft of penis supports two relatively small, rounded cylindrical rami about 200 – 250 µm long; distal part of penis, including rami, bearing numerous short but wide conical setae (Figs 21 D, 25 D). Rest of abdomen vestigial, consisting of two indistinct segments; posterior end of abdomen with shallow cleft (Figs 21 C, 25 C). Remarks: The new species Petrarca rubus is similar to the other three congeners, P. morula, P. goanna and P. nozaaeai, in utilizing Turbinaria corals as hosts. These species are characterized by the large lumpy inflations of the carapace, which are absent in other species of Petrarca. Moreover, P. goanna and P. rubus represent a complex of cryptic species and can only be distinguished by a few ultrastructural details: (1) the presence of numerous large, long conical papillae on the anterioventral part of carapace in P. rubus (only a few in P. goanna) and papillae on the external lateral surface of carapace (absent in P. goanna); (2) the ultrastructure of antennular aesthetasc having three long terminal outgrowths in P. rubus instead of four in P. goanna; and (3) rounded cylindrical rami of penis in P. rubus but squarish rami in P. goanna. Petrarca rubus differs from P. morula and P. nozaaeai in having radial ribs and a crenulated margin of the carapace valves and the presence of bifid teeth on the mandibles. MOLECULAR ANALYSIS	en	Kolbasov, Gregory A., Savchenko, Alexandra S., Yu, Meng-Chen, Tsao, Yao-Feng, Ganmanee, Monthon, Chan, Benny K. K. (2023): Integrative taxonomy, larval biology and functional morphology of the little known gall-forming coral endoparasite Petrarca (Thecostraca: Ascothoracida). Zoological Journal of the Linnean Society 198: 767-801, DOI: 10.1093/zoolinnean/zlad009
