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Neogene and Quaternary ghost shrimps and crabs (Crustacea: Decapoda) from the Philippines

January 2008

January 2008
34:51-76

Authors:

Hiroaki Karasawa

Mizunami Fossil Museum

Hisayoshi Kato

Natural History Museum and Institute, Chiba

Tomoki Kase

National Museum of Nature and Science, Tsukuba

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Twenty species in 17 genera of decapod crustaceans are reported from the Upper Miocene Mapulo Formation in Taysan, Batangas, southern Luzon and the Lower Pleistocene Mandog Formation in Davao City, southeastern Mindanao, Philippines. Of these, Leucosia martini, Nursia bilobata, Pseudophilyra granulimarginata, Cryptolutea warreni, Demania pilipinas, and Hexapus granuliformis are described as new. Neocallichirus dijki (Martin, 1883), new combination is also included. In addition, three new combinations are proposed: Philyra shihchenii for Leucosia shihchenii Hu and Tao, 1979, from the Pliocene of Taiwan, Paranursia acharyai for Nursia acharyai Bachmayer and Mohanti, 1973, from the Miocene of India, and Cryptolutea litoralis for Galene litoralis Collins, Lee, and Noad, 2003, from the Pleistocene of Sarawak.

Maps showing detailed locations of DVO-11 in Davao Province (upper right) and TYS-1 in Batangas Province (lower left).

…

Mandog Formation at DVO-11, abandoned sand quarry in Bolbe, Davao City. A. Columnar section of lowermost part of Mandog Formation exposed at DVO-11. B. Outcrop of upper part of section, showing muddy fine-grained sandstone with concretions. Fossil crabs described in this paper were collected from bedding surface on terrace (ca. 100 m 2 ) above dotted line. C. Photograph showing weathered surface of crabbearing bed. D. Photograph showing articulated specimen in crab-bearing bed.

…

. List of know fossil species of Leucosia. Asterisk indicates extant species.

…

List of fossil species of Philya. Asterisk indicates extant species

…

A, B. Eucrate sp., MGB-CF0024, loc. DVO-11, A, ventral; B, dorsal view. C. Carcinoplax purpurea Rathbun, 1914, MGB-CF0025, loc. DVO-11, carapace, dorsal view. D. Carcinoplax purpurea Rathbun, 1914, MGB-CF0026, loc. DVO-11, carapace, dorsal view. E, F. Carcinoplax purpurea Rathbun, 1914, MGBCF0026, loc. DVO-11, E, dorsal; F, ventral view. G, H. Liagore rubromaculata (de Haan, 1835), MGBCF0017, loc. DVO-11, female, G, ventral; H, dorsal view. I, J. Liagore rubromaculata (de Haan, 1835), MGB-CF0018, loc. DVO-11, male, I, dorsal; J, ventral view. K, L. Liagore rubromaculata (de Haan, 1835), MGB-CF0019, loc. DVO-11, female, K, dorsal; L, ventral view. M, N. Demania sp., MGB-CF0023, loc. DVO-11, carapace, M, frontal; N, dorsal view. Scale bars1 cm.

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Introduction

Decapod fossils are hitherto poorly known

from the Philippines. So far, the only reported

occurrence is the single ghost shrimp, Callianas-

sa dijki Martin, 1883, identiﬁed by Martin (1895)

and Smith (1913) from the Neogene of central

Cebu. In 2003, the National Museum of Nature

and Science, Tokyo (NMNS), and the Mines

and Geosciences Bureau, Philippines (MGB),

launched a joint research project on fossil collec-

tion building and natural history research in the

Philippines. This project aims to establish stan-

dard fossil reference materials in the Philippines,

revive paleontological researches in the MGB,

and understand the origin of marine biodiversity

in the tropical Indo-Western Paciﬁc (see Aguilar

and Kase, 2004). Since its initiation, T. Kase, Y.

M. Aguilar and Y. Kurihara undertook fossil col-

lection surveys in many islands in the Philip-

pines, and discovered a number of unexplored

younger Cenozoic fossil localities. During the

course of paleontological surveys conducted by

NMNS and MGB, a number of decapod fossils

were collected in Taysan, Batangas and Davao

City in Mindanao. A total of 20 species are rep-

resented in the collections, six of which are new

(Table 1). The purpose of this paper is to system-

Neogene and Quaternary Ghost Shrimps and Crabs (Crustacea:

Decapoda) from the Philippines

Hiroaki Karasawa1, Hisayoshi Kato2, Tomoki Kase3, Yolanda Maac-Aguilar4,

Yukito Kurihara3, Hiroki Hayashi5and Kyoko Hagino6

1Mizunami Fossil Museum, Yamanouchi, Akeyo, Mizunami, Gifu 509–6132 Japan

E-mail: GHA06103@nifty.com

2Natural History Museum and Institute, Chiba, Aoba-cho, Chiba 260–8682 Japan

E-mail: katoh@chiba-muse.or.jp

3Department of Geology and Paleontology, National Museum of Nature and Science,

Hyakunin-cho 3–23–1, Shinjyuku, Tokyo 169–0073 Japan

E-mail: kase@kahaku.go.jp; kurihara@kahaku.go.jp

4Petrolab, Mines and Geosciences Bureau, North Ave., Diliman, Quezon City, Philippines

E-mail: yolagui@yahoo.com

5Department of Geoscience, Interdisciplinary Faculty of Science and Engineering, Shimane University,

Shimane 690–8504 Japan

E-mail: hayashi@riko.shimane-u.ac.jp

6Institute for Study of the Earth’s Interior, Okayama University, 827 Yamada, Misasa, Tottori 682–0193 Japan

E-mail: hagino@pheasant.misasa.okayama-u.ac.jp

Abstract Twenty species in 17 genera of decapod crustaceans are reported from the Upper

Miocene Mapulo Formation in Taysan, Batangas, southern Luzon and the Lower Pleistocene Man-

dog Formation in Davao City, southeastern Mindanao, Philippines. Of these, Leucosia martini,

Nursia bilobata, Pseudophilyra granulimarginata, Cryptolutea warreni, Demania pilipinas, and

Hexapus granuliformis are described as new. Neocallichirus dijki (Martin, 1883), new combination

is also included. In addition, three new combinations are proposed: Philyra shihchenii for Leucosia

shihchenii Hu and Tao, 1979, from the Pliocene of Taiwan, Paranursia acharyai for Nursia

acharyai Bachmayer and Mohanti, 1973, from the Miocene of India, and Cryptolutea litoralis for

Galene litoralis Collins, Lee, and Noad, 2003, from the Pleistocene of Sarawak.

Key words : Neogene, Crustacea, Decapoda, Thalassinidea, Brachyura, Philippines.

Bull. Natl. Mus. Nat. Sci., Ser. C, 34, pp. 51–76, December 22, 2008

atically document these decapod species.

The ﬁrst and second authors are responsible

for the taxonomy of the crabs, while the others

are for the description of geology and collecting

sites. The specimens described in this paper are

stored in the MGB and NMNS with preﬁx MGB

and NMNS, respectively.

Description of Fossil Localities

By T. Kase, Y. M. Aguilar, Y. Kurihara, H.

Hayashi and K. Hagino

The fossil decapods described in this paper

were collected from two sites: Locality TYS-1

(Mapulo Formation) and locality DVO-11 (Man-

dog Formation). Details of the localities are de-

scribed below.

Locality TYS-1

Location: Limestone quarry at Fortune Ce-

ment Co. Ltd., Barangay Mapulo, Taysan munici-

pality in Batangas Province, southern Luzon

(13°44.472!N, 121°11.240!E) (Fig. 1).

Stratigraphy: Locality TYS-1 is from the lime-

stone quarry located in Mapulo Hill, Taysan that

forms a small topographic high within the Pleis-

tocene volcanic sediments. The sedimentary beds

exposed in this quarry consist of an upper ca. 30-

m-thick coralline limestone and a lower ca. 50-

m-thick, slightly consolidated dark-gray muddy

sandstone. The upper limestone facies was as-

signed to the Mapulo Limestone by Avila (1980)

and was accordingly adopted by the Philippine

Bureau of Mines and Geosciences (1985). Kase

and Aguilar (2004) elevated the Mapulo to for-

mational level to include the upper limestone and

lower sandstone beds. Fossil mollusks are quite

common in the upper part of the lower muddy

sandstone beds. In a preliminary examination,

Kase and Aguilar (2004) identiﬁed a total of 55

molluscan species, 36 of which are identical to

modern species that are mostly dwellers in subti-

dal to bottoms of about 50 m in depth. Fossil

crabs are rather sporadic and found in ﬂoated cal-

careous concretions.

Age: Philippine Bureau of Mines and Geo-

sciences (1985) assigned the Mapulo Limestone

52 H. Karasawa et al.

Table1. List of species from the Mapulo and Mandog Formation

Collecting site, age and formation

Species TYS-1 Late Miocene DVO-11 Pleistocene

Mapulo Formation Mandog Formaiton

Callianassa (s. l.) sp. "

Neocallichirus dijki (Martin, 1883), new combination "

Neocallichirus sp. ""

Raninoides sp. "

Calappa pustulosa Alcock, 1896 "

Calappa sp. "

Leucosia martini Karasawa & Kato, new species "

Nursia bilobata Karasawa & Kato, new species "

Philyra sp. cfr. P. actidens Chen, 1987 "

Pseudophilyra granulimarginata Karasawa & Kato, new species "

Galene bispinosa (Herbst, 1783) "

Cryptolutea warreni Karasawa & Kato, new species "

Liagore rubromaculata (de Haan, 1835) "

Demania pilipinas Karasawa & Kato, new species "

Demania sp. "

Eucrate sp. "

Carcinoplax purpurea Rathbun, 1914 "

Hexapus granuliformis Karasawa & Kato, new species "

Podophthalmus vigil (Fabricius, 1798) "

Pinnixa sp. "

a late Miocene to Pliocene age. This was sub-

stantiated by nannofossil analysis made by H.

Kameo of Chiba University on the sandstone

sample that can be assigned to nannofossil zone

CN9 (8.36 to 5.56 Ma), equivalent to a late Late

Miocene age.

Locality DVO-11

Location: Abandoned sand quarry at Bolbe,

Davao City, southeastern Mindanao (7°04!01#N,

125°34!05#E) (Fig. 1).

Stratigraphy: Younger Cenozoic, ﬂuvio-deltatic

to shallow marine sedimentary sequences in the

area around Davao City, southeastern Mindanao

consist of lower Masuhi and upper Mandog for-

mations which are overlain unconformably by the

younger Pleistocene Apo Volcanics and the

Samal Limestone. According to the geological

map published by Mines and Geosciences Bu-

reau, Philippines (Philippine Bureau of Mines

and Geoscience, 1984), DVO-11 is within the

Mandog Formation that is referred vaguely to

early Pleistocene in age. The Masuhi Formation

in this area is represented by deltaic conglomer-

ate of more than 100 m thick, while the overlying

Mandog Formation represents a transgressive fa-

cies succession that consists of sandstone, marl

and coralline limestone. The sandstone beds at

DVO-11 is the lowermost part of the Mandog

Formation, remarkably fossiliferous, and contain

Neogene and Quaternary Crustaceans from Philippines 53

Fig. 1. Maps showing detailed locations of DVO-11 in Davao Province (upper right) and TYS-1 in Batangas

Province (lower left).

diverse mollusks that are mostly identical to

modern species found today in bottoms of open

marine, around 50 m in depth (Bed c in Fig. 2A).

The fossil crabs described here are from an upper

horizon of the beds exposed in this quarry (Bed a

in Fig. 2A). The crab-bearing beds are composed

of ﬁne-grained sandstone with abundant calcare-

ous concretions that may have originated by bur-

rows of benthic animals (Fig. 2B–D). The crabs

are abundant in a weathering bedding surface

(Fig. 2B), and often articulated in good condition

(Fig. 2D).

Age: We analyzed planktonic foraminifers and

nannofossils from Bed b just below the crab-

bearing bed (Fig. 2A). Bed b yielded beautiful-

ly preserved planktonic foraminifer abundant-

ly.

Species identiﬁed include Globigerina bul-

loides, G. decoraperta, Globoturborotalita cf.

rubescens, Globigerinella sp., Globigerinita

glutinata, Globigerinoides bulloideus, G. rubber,

G. sacculifer, Globorotalia cf. crassaformis, G.

menardii, Neogloboquadrina humerosa s.l., Pul-

leniatina obliquiloculata (dextrally coiled form),

P. obliquiloculata (sinistrally coiled form), Orbu-

lina spp. and some others. These species are not

diagnostic in determining detailed age, except

that the abundance occurrence of dextral form of

Pulleniatia obliquiloculata suggests an age

younger than 3.95 Ma (Berggren et al., 1995).

Bed b yielded moderate to well-preserved Pleis-

tocene calcareous nannofossils as well as re-

worked specimens from older Pliocene sedimen-

tary rocks. The calcareous nannofossil species

identiﬁed from this sample are; Calcidiscus lep-

toporus, Gephyrocapsa caribbeanica, medium-

sized Gephyrocapsa oceanica (3.5–5.0

m in

length of placolith), Discoaster broweri, Spheno-

lithus sp., and Reticulofenestra minutula. The oc-

currence of medium-sized G. oceanica indicates

that the sample is younger than 1.7Ma (Rafﬁ et

al., 2006). Therefore, these micropaleontological

data demonstrate that Bed b is broadly assigned

54 H. Karasawa et al.

Fig. 2. Mandog Formation at DVO-11, abandoned sand quarry in Bolbe, Davao City. A. Columnar section of

lowermost part of Mandog Formation exposed at DVO-11. B. Outcrop of upper part of section, showing

muddy ﬁne-grained sandstone with concretions. Fossil crabs described in this paper were collected from bed-

ding surface on terrace (ca. 100 m2) above dotted line. C. Photograph showing weathered surface of crab-

bearing bed. D. Photograph showing articulated specimen in crab-bearing bed.

to the Pleistocene. Taking the stratigraphic posi-

tion into consideration, however, the age of the

crab-bearing beds in the lowest part of the Man-

dog Formation is estimated to be early Pleis-

tocene.

Systematics

By H. Karasawa and H. Kato

Infraorder Thalassinidea Latreille, 1831

Superfamily Callianassoidea Dana, 1852

Family Callianassidae Dana, 1852

Subfamily Callianassinae Dana, 1852

Genus Callianassa (s. l.) Leach, 1814

Type species: Cancer (Astacus) subterraneus

Montagu, 1808, by monotypy.

Callianassa (s. l.) sp.

(Fig. 3A)

Material examined: MGB-CF0001 and NMNS

PA16389 from TYS-1.

Remarks: The present specimens are the poor-

ly preserved propodi of the major chelipeds.

Therefore, the generic placement of the species

awaits the discovery of more well-preserved

material and it is considered best to place the

species in Callianassa (s. l.). The smooth lateral

surface and dorsal and ventral margins easily dis-

tinguish it from the following two callichirhines,

Neocallichirus dijki (Martin, 1883) and Neocal-

lichirus sp.

Subfamily Callichirinae Manning and Felder,

1991

Genus Neocallichirus Sakai, 1988

Type species: Neocallichirus horneri Sakai,

1988, by original designation.

Included fossil species: see Schweitzer et al.

(2008).

Remarks: Most recently, Schweitzer et al.

(2006b, 2008) reviewed the fossil species within

Neocallichirus and included 16 species in the

genus. Among these, Neocallichirus grandis

Karasawa and Goda, 1996, has been transferred

to Grynaminna Poore, 2000, based upon the

characters of the chelipeds (Obata and Hayashi,

2000). After that, Sakai (2005) synonymised

Grynaminna with Podocallichirus Sakai, 1999;

therefore, Karasawa et al. (2006) removed Gry-

naminna grandis to Podocallichirus. Additional-

ly, Callianassa dijki Martin, 1883, is herein

moved to the present genus.

Neocallichirus dijki (Martin, 1883),

new combination

(Fig. 3B, C)

Callianassa dijki Martin, 1883, 36, pl. 3, ﬁgs.

31–33.

Material examined: MGB-CF0002 and NMNS

PA16390 from TYS-1.

Emended diagnosis: Large-sized major che-

liped. Palm about as long as high; lateral and

mesial surfaces coarsely granulate distally; ven-

tral margin denticulate; dorsal margin ﬁnely den-

ticulate; distal margin granulated. Fixed ﬁnger

slightly shorter than palm, gently curved dorsally,

occlusal margin ﬁnely serrated. Dactylus coarse-

ly granulated proximally; ventrolateral surface

pitted; tip acutely pointed, hooking ventrally; oc-

clusal margin bearing large denticles varying in

size.

Remarks: Martin (1883) described Callianassa

dijki from the Miocene of Java. After that, Martin

(1895) and Smith (1913) recorded this species

from the Miocene deposits of Philippines. Cal-

lianassa dijki possesses characters of the propo-

dus of the major cheliped most like those of the

extant Neocallichirus karumba (Poore and Grif-

ﬁn, 1979), a senior synonym of Neocallichirus

kempi Sakai, 1999 (see Dworschak, 2008), from

East Asia, Australia, and India, previously as-

signed to Callianassa maxima A. Milne Ed-

wards, 1870; therefore, this species is moved to

Neocallichirus.

Neogene and Quaternary Crustaceans from Philippines 55

56 H. Karasawa et al.

Neocallichirus sp.

(Fig. 3Q, R)

Material examined: MGB-CF0003 from DVO-

11; NMNS PA16391 from TYS-1.

Remarks: This species is represented by a bro-

ken propodus of the larger cheliped. A large-

sized propodus with serrated dorsal, ventral, and

distal margins and a granulated dorso-mesial sur-

face of the ﬁxed ﬁnger characterize it. A smooth

lateral surface readily distinguishes this species

from Neocallichirus dijki. This species is similar

to Neocallichirus sp. from the Pliocene Shimajiri

Group, reported by Karasawa (1997), but differs

in having a serrated dorsal margin and a granu-

lated dorso-lateral surface of the ﬁxed ﬁnger.

Infraorder Brachyura Latreille, 1802

Section Raninoida de Haan, 1841

Superfamily Raninoidea de Haan, 1841

Family Raninidae de Haan, 1841

Subfamily Raninoidinae Lo”renthey in Lo”renthey

and Beurlen, 1929

Genus Raninoides H. Milne Edwards, 1837

Type species: Raninoides laevis Latreille,

1825, by monotypy.

Included fossil species: see Schweitzer et al.

(2006a).

Remarks: Although Schweitzer et al. (2006a)

made a list of the known extant and fossil species

within the genus, there is an additional species,

Raninoides fulungensis Hu and Tao, 2000, from

the Oligocene–Miocene of Taiwan.

Raninoides sp.

(Fig. 3S)

Material examined: MGB-CF0004 and NMNS

PA16392 from DVO-11.

Remarks: In the present specimen, detailed

characters of the fronto-orbital region is wanting.

The speciﬁc identiﬁcation of this species awaits

the discovery of more well-preserved specimens.

Section Eubrachyura de Saint Laurent, 1980

Superfamily Calappoidea de Haan, 1833

Family Calappidae de Haan, 1833

Genus Calappa Weber, 1795

Type species: Cancer granulatus Linnaeus,

1758, by subsequent designation of Latreille

(1810).

Included fossil species: see Schweitzer et al.

(2006b).

Remarks: Schweitzer et al. (2006b) reviewed

the fossil species of Calappa and recognized 19

species in the fossil record. In their review, three

species lack: two extinct, Calappa chungii Hu

and Tao, 1984, and Calappa oboui Hu and Tao,

1996, from the Pliocene of Taiwan; and one ex-

tant Calappa pustulosa Alcock, 1896, from the

Pliocene of Taiwan (Hu and Tao, 2000; 2004).

Neogene and Quaternary Crustaceans from Philippines 57

Fig. 3. A. Callianassa (s. l.) sp., MGB-CF0001, loc. TYS-1, propodus of major cheliped, lateral view. B, C.

Neocallichirus dijki (Martin, 1883), new combination, MGB-CF0002, loc. TYS-1, propodus of major che-

liped, B, lateral; C, mesial view. D, E. Nursia bilobata Karasawa and Kato, new species, MGB-CF0009

(holotype), loc. TYS-1, carapace, D, dorsal; E, ventral view. F, G. Galene bispinosa (Herbst, 1783), MGB-

CF0012, loc. TYS-1, carapace, F, dorsal; G, ventral view. H-J. Pseudophilyra granulimarginata Karasawa

and Kato, new species, MGB-CF0011 (holotype), loc. TYS-1, carapace, H, dorsal; I, ventral; J, lateral view.

K. Calappa sp., MGB-CF0006, loc. DVO-11, carapace, dorsal view. L, M. Leucosia martini Karasawa and

Kato, new species, MGB-CF0008 (paratype), loc. DVO-11, carapace, L, dorsal; M, lateral view. N, O. Leu-

cosia martini Karasawa and Kato, new species, MGB-CF0007 (holotype), loc. DVO-11, carapace, N, lateral;

O, dorsal view. P. Philyra sp. cfr. P. actidens Chen, 1987, MGB-CF0010, loc. TYS-1, carapace, dorsal view.

Q, R. Neocallichirus sp., MGB-CF0003, loc. DVO-11, propodus of major cheliped, Q, mesial; R, lateral

view. S. Raninoides sp., MGB-CF0004, loc. DVO-11, carapace, dorsal view. T. Calappa pustulosa Alcock,

1896, MGB-CF0005, loc. DVO-11, carapace, dorsal view. Scale bars$1cm.

Calappa pustulosa Alcock, 1896

(Fig. 3T)

Material examined: MGB-CF0005 from DVO-

11.

Remarks: The dorsal surface of the present

specimen is rather smooth because of erosion.

This species is also recorded from the Pliocene

Shimajiri Group of the Miyako island, Ryukyus,

Japan (Karasawa and Nobuhara, 2008). More

complete material will be necessary to conﬁrm

identiﬁcation of this species.

Calappa sp.

(Fig. 3K)

Material examined: MGB-CF0006 and NMNS

PA16393 from DVO-11.

Remarks: The specimens are a poorly pre-

served carapace and cheliped. However, the re-

maining carapace characters may be similar to

the extant Calappa clyeata (Borradaile, 1903).

Superfamily Leucosioidea Samouelle, 1819

Family Leucosiidae Samouelle, 1819

Genus Leucosia Weber, 1795

Type species: Cancer craniolaris Linnaeus,

1758, by subsequent designation of Holthuis

(1959).

Included fossil species: Table 2.

Remarks: Most recently, Galil (2003a, b,

2005a, b, 2006) reviewed Leucosia and erected

ﬁve new genera, Euclosia Galil, 2003b, Ur-

nalana Galil, 2005a, Seulocia Galil, 2005b,

Coleusia Galil, 2006a, and Soceulia Galil,

2006b, all which were previously assigned to the

species of Leucosia. In her revision, Leucosia

was restricted to four species, L. craniolaris (Lin-

naeus, 1758), L. moresbiensis Haswell, 1880, L.

punctata Bell, 1855, and L. rubripalma Galil,

2003a, and most species were transferred to her

new genera. Her work is continuous and the re-

maining species have not yet been assigned to

any genus.

Eighteen species of Leucosia sensu lato have

58 H. Karasawa et al.

Table2.List of know fossil species of Leucosia. Asterisk indicates extant species.

Original status Current status Age and locality References

L. anatum (Herbst, 1783)* Leucosia s. l. M. Pleistocene, Japan Karasawa & Tanaka (1994);

Kato & Karasawa (1997)

L. calcarata Collins et al., 2003 Leucosia s .l. E. Miocene, Sarawak Collins et al. (2003)

L. craniolaris (Linnaeus, 1758)* Leucosia s. s. Pliocene, Taiwan Hu & Tao (1996)

L. formosensis Sakai, 1937 Philyra Pleistocene, Taiwan Hu & Tao (1996)

L. haematsticta Adams & White, 1848* Urnalana M. Pleistocene, Japan Karasawa & Goda (1996);

parahaematsticta Kato & Karasawa (1998);

Galil, 2005 Kobayashi et al. (2008)

L. longiangulata Morris & Collins, 1991 Leucosia s .l. Pliocene, Brunei Morris & Collins (1991)

L. martini Karasawa & Kato, new species Leucosia s. l. Pleistocene, Philippines This paper

L. obtusifrons de Haan, 1841 Euclosia Pliocene, Taiwan Hu & Tao (1996)

Neogene, Java Martin (1879)

L. ovalata Hu & Tao, 1996 Leucosia s. l. L. Miocene, Taiwan Hu & Tao (1996)

L. rhomboidalis de Haan, 1841* Seulocia L. Pliocene–Pleistocene Hu & Tao (1996)

L. serenei Morris & Collins, 1991 Leucosia s. l. Pliocene, Brunei Morris & Collins (1991)

L. shihcheni Hu & Tao, 1979 Philyra Pliocene, Taiwan Hu & Tao, 1979; This work

L. subrhomboidalis Desmarest, 1822 Leucosia s. l. Pleistocene, Indo-W. Paciﬁc Glaessner (1929)

L. subrhomboidea Hu & Tao, 1996 Leucosia s. l. L. Miocene, Taiwan Hu & Tao (1996)

L. taiwanica Hu & Tao, 1996 Leucosia s. l. E. Pleistocene, Taiwan Hu & Tao (1996)

L. takamii Karasawa, 1993 Leucosia s. l. Pleistocene, Japan Karasawa (1993)

L. tricarinata Martin, 1880 Leucosia s. l. Miocene, Java Martin (1880)

L. tutongensis Morris & Collins, 1991 Leucosia s. l. Pliocene, Brunei Morris & Collins (1991)

L. unidentata de Haan, 1841* Euclosia Pliocene, Taiwan Hu & Tao (1996)

Pleistocene, Japan Karasawa (2000)

Miocene, Java Martin (1879)

been known in the fossil record. However, it is

very difﬁcult to adapt Galil’s classiﬁcation for the

extinct species because of lacking detailed char-

acters of the gonopod 1 of males within the fos-

sils. Therefore, it is considered best to place the

extinct species within Leucosia sensu lato for the

time being. Among these fossils, Leucosia shi-

hchenii Hu and Tao, 1979, from the Pliocene of

Taiwan, has a granulated dorsal carapace, a well-

developed subhepatic facet, and well-deﬁned car-

diac and intestinal regions; therefore, this species

should be assigned to Philyra Leach, 1817. Leu-

cosia formosensis Sakai, 1937, described from

the Pleistocene of Taiwan by Hu and Tao (1996),

has deep cervical and branchiocardiac grooves

and a well-deﬁned intestinal region. These speci-

mens are identical with the carapace of Philyra,

but its species-level identiﬁcation awaits the dis-

covery of more well-preserved specimens.

Leucosia martini Karasawa and Kato,

new species

(Fig. 3L–O)

Material examined: MGB-CF0007 (holotype),

MGB-CF0008 (paratype), and NMNS PA16394

from DVO-11.

Diagnosis: Carapace elongate rhomboidal,

widest a little posterior to mid-length. Front

narrow, strongly projected anteriorly, upturned

dorsally. Anterolateral margin slightly sinuous,

granulate, rimmed. Epibranchial angle not pro-

nounced. Posterolateral margin granulate, rimed.

Posterior margin gently convex, granulated,

rimmed. Epimeral edge invisible dorsally, ﬁnely

granulate. Thoracic sinus deep; anterior edge not

investigated, deﬁned by nearly straight, obtusely

granulated edge of pterygostomian region; three

rows of granules present above coxa of cheliped.

Etymology: After K. Martin, who ﬁrst reported

fossil decapods from the Philippines.

Description: Carapace elongate rhomboidal in

outline, width about 85% carapace length, widest

a little posterior to mid-length. Fronto-orbital

margin narrow, about 15% carapace width. Dor-

sal surface glabrous, smooth, regions not deﬁned.

Front strongly projected anteriorly, upturned dor-

sally. Frontal margin weakly trilobed. Orbit

small. Anterolateral margin slightly sinuous,

granulate, rimmed. Epibranchial angle not pro-

nounced. Posterolateral margin also granulate,

rimmed. Posterior margin gently convex, granu-

lated, rimmed, about 25% carapace width.

Epimeral edge invisible dorsally, ﬁnely granulate,

joining posterior margin. Thoracic sinus deep;

anterior edge not investigated, deﬁned by nearly

straight, obtusely granulated edge of pterygosto-

mian region; three rows of granules present

above coxa of cheliped; granules on upper row

largest, nearly close to granules on middle row;

lower row consisting of ﬁne granules.

Remarks: The present new species possesses

characters of the thoracic sinus of the carapace

most like those of Leucosia tricarinata Martin,

1880, but differs in having three granulated rows

above the coxa of cheliped in the thoracic sinus

(vs. two granulated rows in L. tricarinata).

Genus Nursia Leach, 1817

Type species: Nursia hardwickii Leach, 1817,

by monotypy.

Included fossil species: Nursia bilobata, new

species; N. sp. aff. N. japonica Sakai, 1935 (Kato

and Karasawa, 1998) (extant?).

Remarks: The genus Nursia is recognized as a

heterogeneous group (Ihele, 1918; Serène and

Soh, 1976; Komatsu and Takeda, 2003); there-

fore, Serène and Soh (1976) erected a new genus

Paranursia for N. abbreviata Bell, 1855, and Ko-

matsu and Takeda (2003) proposed a new genus

Nobiliella for N. jousseaumei Nobili, 1905 and

N. jousseaumei var. cornigera Nobili, 1905.

However, a generic level reconsideration of re-

maining species of Nursia is needed (Serène and

Soh, 1976; Komatsu and Takeda, 2003). Nursia

acharyai Bachmayer and Mohanti, 1973, is

known from the Miocene of India. This species is

represented by a single specimen of an eroded

carapace. In Nursia acharyai the dorsal carapace

has a median longitudinal ridge and oblique

Neogene and Quaternary Crustaceans from Philippines 59

ridges running from the mesogastric region

across the epibranchial regions and lacks hepatic

ridges, and the posterior margin bears no lobe.

Those are deﬁnitive characters of Pa ranursia

(Serène and Soh, 1976; Poore, 2004); therefore,

N. acharyai is moved to Paranursia. Pa ranursia

acharyai differs from P. abbreviata by having

dentate anterolateral margins and concave pos-

terolateral margins.

Nursia bilobata Karasawa and Kato,

new species

(Fig. 3D, E)

Material examined: MGB-CF0009 (holotype)

from TYS-1.

Diagnosis: Carapace rhomboidal, widest about

at mid-length. Pterygostomian margin well de-

veloped, prominent medially. Anterolateral mar-

gin bearing 3 teeth; anterior two teeth low; last

one projected, directed posterolaterally. Postero-

lateral margin with broadly triangular tooth at

posterior third. Posterior margin with 2 triangular

lobes. Dorsal surface strongly convex; elevated

regions covered with ﬁne granules and tubercles.

Weak median ridge present on anterior mesogas-

tric process. Mesogastric region bearing pair of

large tubercles, with weak, oblique ridge extend-

ing from anterolateral margin across hepatic re-

gion. Arcuate ridge running from anterior part of

inverted-triangle cardiac elevation to epi-

branchial ridge, extending from hepatic ridge to

last anterolateral tooth. Strong transverse ridge

present on intestinal and metabranchial regions.

Etymology: The trivial name refers to the bilo-

bate posterior margin.

Description: Carapace rhomboidal in outline,

appears to be slightly longer than wide, widest

about at mid-length. Fronto-orbital margin about

28% carapace width, detailed characters wanting.

Pterygostomian margin well developed, elevated

medially. Anterolateral margin bearing 3 teeth;

anterior tooth low, broad, directed anterolaterally,

behind posterior end of pterygostomian margin;

second one low, broadly triangular, directed later-

ally; last one projected, directed posterolaterally.

Posterolateral margin bearing posterolaterally di-

rected, broadly triangular tooth at posterior third.

Posterior margin bilobate, slightly shorter than

fronto-orbital margin; lobes triangular with

rounded tip. Dorsal surface strongly convex lon-

gitudinally and transversely; elevated regions

covered with ﬁne granules and tubercles. Weak

median ridge present on anterior mesogastric

process. Mesogastric region bearing pair of large

tubercles, with low, oblique ridge extending from

anterolateral margin across hepatic region. Car-

diac region most convex; arcuate ridge well de-

ﬁned, running from anterior part of inverted-tri-

angle cardiac elevation to epibranchial ridge.

Epibranchial ridge strong, extending from hepat-

ic ridge to last anterolateral tooth. Strong trans-

verse ridge extending from tooth on posterolater-

al margin across intestinal and metabranchial re-

gions.

Remarks: The present new species is most

similar to the extant Nursia nasuta Alcock, 1896,

because the carapace is relatively narrow with a

weak hepatic ridge on the dorsal region, and the

posterior margin consists of two lobes. However,

the present species differs from N. nasuta in that

a well deﬁned arcuate ridge extends from the an-

terior part of the cardiac region to the epi-

branchial ridge, the intestinal and mesobranchial

regions are much longer than those of N. nasuta,

and two posterior lobes are well developed and

are more strongly protruded posteriorly than in

N. nasuta.

Genus Philyra Leach, 1817

Type species: Leucosia globus Fabricius, 1775,

by subsequent designation of H. Milne Edwards

(1837).

Included fossil species: Philyra alveola Hu and

Tao, 2000; P. cranium (Desmarest, 1822); P. fer-

rica Hu and Tao, 1996; P. granulosa Morris and

Collins, 1991; P. hayasakai Karasawa and Inoue,

1992; P. heterograna Ortmann, 1892 (also ex-

tant); P. miyamotoi Karasawa and Kishimoto,

1996; P. nishimotoi Karasawa, 1989; P. pisum de

60 H. Karasawa et al.

Haan, 1841 (also extant); P. plana Karasawa,

1989; P. platycheir de Haan, 1841 (also extant);

P. scabriuscula (Fabricius, 1793) (also extant); P.

shihcheni (Hu and Tao, 1979), new combination;

P. syndactyla Ortmann, 1892 (also extant); P.

tanakai Karasawa, 1993; P. tridentata Karasawa,

1993; P. trusanensis Collins et al., 2003; Philyra

sp. cfr. P. actidens Chen, 1987.

Remarks: The systematic status of Philyra shi-

hcheni (Hu and Tao, 1979) was discussed above.

The stratigraphic and geographic distribution of

the known fossils of Philyra is summarized in

Table 3.

Philyra sp. cfr. P. actidens Chen, 1987

(Fig. 3P)

Material examined: MGB-CF0010 from TYS-

Remarks: The present specimen is incomplete

but bears similarity to the extant P. actidens from

the East China Sea, including the dorsal carapace

densely covered with tubercles. We refer the

specimen to the species provisionally because it

lacks the maxilliped 3 and male gonopod 1,

which are considered very important for the

identiﬁcation of the genus.

Genus Pseudophilyra Miers, 1879

Type species: Pseudophilyra tridentata Miers,

1879, by subsequent designation of Rathbun

(1922).

Included fossil species: Pseudophilyra elon-

gatella Hu and Tao, 1996.

Pseudophilyra granulimarginata Karasawa

and Kato, new species

(Fig. 3H–J)

Material examined: MGB-CF0011 (holotype)

from TYS-1.

Diagnosis: Carapace elongate rhomboidal,

widest at mid-length. Fronto-orbital margin nar-

row, about 15% carapace width. Dorsal surface

smooth; regions not deﬁned. Anterolateral mar-

gin slightly sinuous, granulate, rimmed. Epi-

branchial angle not pronounced. Posterolateral

margin convex, granulate, rimmed. Posterior

margin nearly straight, granulated, rimmed.

Etymology: The trivial name refers to the gran-

Neogene and Quaternary Crustaceans from Philippines 61

Table3 List of fossil species of Philya. Asterisk indicates extant species

Species Age and locality References

P. alveola Hu & Tao, 2000 Pliocene, Taiwan Hu & Tao (2000)

P. cranium (Desmarest, 1822) Pleistocene?, Indo-Paciﬁc Glaessner (1929)

P. ferrica Hu & Tao, 1996 U. Miocene, Taiwan Hu & Tao (1996)

P. granulosa Morris & Collins, 1991 M. Miocene, Sarawak Morris & Collins (1991)

P. hayasakai Karasawa & Inoue, 1992 M. Miocene, Japan Karasawa & Inoue (1992)

P. heterograna Ortmann, 1892* Pleistocene, Japan Kato & Koizumi (1992)

P. miyamotoi Karasawa & Kishimoto, 1996 M. Miocene, Japan Karasawa & Kishimoto (1996)

P. nishimotoi Karasawa, 1989 E. Miocene, Japan Karasawa (1989)

P. pisum de Haan, 1841* Pleistocene, Japan Kato & Koizumi (1992)

P. plana Karasawa, 1989 M. Miocene Karasawa (1989)

P. platycheir de Haan, 1841* Pliocene, Taiwan Hu & Tao (2000)

Pliocene–Pleistocene, Japan Karasawa (1993),

Karasawa & Tanaka (1994)

P. scabriuscula (Fabricius, 1793)* Pliocene, Java, Sumatra Van Straelen (1938)

P. shihcheni (Hu & Tao, 1979) M. Miocene, Taiwan Hu & Tao (1979)

P. syndactyla Ortmann, 1892* Pleistocene, Japan Kato & Koizumi (1992),

Kato & Karasawa (1998),

Kobayashi et al. (2008)

P. tanakai Karasawa, 1993 Pliocene, Japan Karasawa (1993)

P. tridentata Karasawa, 1993 Pliocene, Japan Karasawa (1993)

ulated anterolateral, posterolateral, and posterior

margins.

Description: Carapace elongate rhomboidal in

outline, width about 87% carapace length, widest

at mid-length. Fronto-orbital margin narrow,

about 15% carapace width. Dorsal surface

glabrous, smooth, regions not deﬁned. Front pro-

jected anteriorly, upturned dorsally. Frontal mar-

gin weakly trilobed. Orbit small. Anterolateral

margin slightly sinuous, granulate, rimmed. Epi-

branchial angle not pronounced. Posterolateral

margin convex, granulate, rimmed. Posterior

margin nearly straight, granulated, rimmed, about

23% carapace width. Epimeral edge invisible

dorsally, ﬁnely granulate, joining posterior mar-

gin. Thoracic sternites 1–3 completely fused,

much wider than long; sternite 4 longest of all

sternites, lateral margin deeply notched behind

coxa of cheliped; sulcus between sternites 3 and

4 medially interrupted by narrow, triangular ster-

no-abdominal cavity; sulci between sternites 4/5,

5/6, and 6/7 incomplete medially. Episternites

4–7 narrow, granulated ventrally, protruded pos-

teriorly, separated from each sternite by rather

deep grooves; episternite 8 small, granulate. Ster-

no-abdominal cavity deep; anterior end reaching

sternites 2 and 3.

Remarks: The present new species is most

similar to Pseudophilyra sp., an unnamed species

described from the Recent Philippines by Tan

(1996), but differs in having a narrow fronto-or-

bital margin and a convex posterolateral margin.

This species is easily distinguished from the ex-

tinct P. elongatella from the upper Miocene of

Taiwan by having a narrow fronto-orbital margin

and granulated anterolateral, posterolateral, and

posterior margins. In the carapace characters, P.

granulimarginata may be similar to Leucosia

martini, but differs in lacking the thoracic sinus.

Superfamily Xanthoidea MacLeay, 1838, sensu

Karasawa and Schweitzer, 2006

Family Pilumnidae Samouelle, 1819, sensu

Karasawa and Schweitzer, 2006

Subfamily Galeninae Alcock, 1898

Genus Galene de Haan, 1833

Type species: Cancer bispinosus Herbst, 1783,

by monotypy.

Included fossil species: G. bispinosa (Herbst,

1783) ($G. hainanesis Hu and Tao, 1979) (also

extant); G. granulifera Lin, 1947; G. obscura A.

Milne Edwards, 1865; G. stipata Morris and

Collins, 1991.

Remarks: Hu and Tao (1979) described a new

species, Galene hainanesis, based upon nine

specimens collected from the Hainan island,

China (Hu and Tao, 1979; p. 148). These speci-

mens appear to be brought from drugstores of

Taipei, Taiwan, and they showed that they oc-

curred in the Pleistocene deposits (Hu and Tao,

1979; p. 148). Comparison of the descriptions

and illustrations of G. hainanesis and G.

bispinosa indicates that they are very similar in

carapace and pereiopod morphology. Therefore,

Galene hainanesis should be synonymised with

G. bispinosa. Additionally, examination of the il-

lustrated specimen (Hu and Tao, 1979, pl. 1, ﬁgs.

3, 4; registered number, MTNU2002, deposited

in the Museum of Taiwan Normal University) of

G. hainanesis suggests that it is not referable to

that species. Hu and Tao (1979, p. 148) stated,

“This small sized skeleton is possibly an imma-

ture individual”. However, in the specimen, the

dorsal surface is smooth, the anterolateral margin

is entire without anterolateral teeth, the thoracic

sternum is very wide without deep lateral

grooves on sternite 4, the telson of male ab-

domen is triangular and not elongate, and the lat-

eral surface of the cheliped is smooth with a

keeled ventro-lateral margin. These characters

lack in Galene and are observed in the rhizopine

genus Arges de Haan, 1835. However, detailed

examination of the material will be necessary to

conﬁrm the taxonomic status of it. Two other fos-

sil species were previously assigned to the genus,

but G. proavita Glaessner, 1960, was moved to

Carcinoplax H. Milne Edwards, 1852 (Karasawa

and Kato, 2003) and G. litoralis Collins, Lee, and

Noad, 2003, is transferred to the rhizopine genus

Cryptolutea Ward, 1936, discussed below.

The genus Galene has been recorded from the

62 H. Karasawa et al.

Neogene and Quaternary Crustaceans from Philippines 63

Fig. 4. A, B. Cryptplutea warreni Karasawa and Kato, new species, MGB-CF0015 (holotype), loc. DVO-11,

carapace and chelipeds, A, frontal; B, dorsal view. C, D. Cryptplutea warreni Karasawa and Kato, new

species, MGB-CF0016 (paratype), loc. DVO-11, carapace, C, frontal; D, dorsal view. E, F. Galene bispinosa

(Herbst, 1783), MGB-CF0012, loc. TYS-1, carapace and cheliped, E, fronral; F, dorsal view. G. Galene

bispinosa (Herbst, 1783), MGB-CF0014, loc. TYS-1, propodus of right cheliped, lateral view. H-J, Demania

pilipinas Karasawa and Kato, new species, MGB-CF0021 (holotype), loc. DVO-11, carapace, chelipeds, and

pereiopods, H, frontal; I, ventral; J, dorsal view. K, Demania pilipinas Karasawa and Kato, new species,

MGB-CF0022 (paratype), loc. DVO-11, carapace, chelipeds, and pereiopods, dorsal view. Scale bars$1cm.

fossil record from the Miocene of Taiwan (Lin,

1947; Hu and Tao, 1979; Hu and Tao, 1996); the

Pliocene of Taiwan (Hu and Tao, 1996), Philip-

pines (this work), and Brunei (Morris and

Collins, 1991); the Pleistocene of Java (Böhm,

1922), and the Holocene of Australia (Etheridge

and McCulloch, 1916).

Galene bispinosa (Herbst, 1783)

(Figs. 3F, G; 4E–G)

Material examined: MGB-CF0012, MGB-

CF0013, MGB-CF0014, and NMNS PA16395

from TYS-1.

Remarks: The fossil occurrence of this species

has previously been reported from the Pleis-

tocene of Taiwan (Hu and Tao, 1996) and the

Holocene of Australia (Etheridge and McCul-

loch, 1916).

Subfamily Rhizopinae Stimpson, 1858

Genus Cryptolutea Ward, 1936

Type species: Cryptolutea lindemanensis Ward,

1936, by monotypy.

Included fossil species: Cryptolutea litoralis

(Collins, Lee, and Noad, 2003), new combina-

tion; C. warreni, new species.

Remarks: Collins et al. (2003) described the

new galenine species, Galene litoralis Collins et

al., 2003, from the Pleistocene of Sarawak. How-

ever, G. litoralis is moved to Cryptolutea because

the carapace is rounded rectangular in outline

and the anterolateral margins consist of three

lobes. Galene, in contrast with Cryptolutea, has a

transversely hexagonal carapace with three an-

terolateral spines. Additionally, Galene bears

more or less deﬁned metabranchial regions,

which Cryptolutea lack. Cryptolutea litoralis re-

sembles the extant Cryptolutea sagamiensis

(Sakai, 1935), but differs in having well deﬁned

mesogastric and cardiac regions and well devel-

oped anterolateral lobes.

The Rhizopinae comprises 22 genera within

the present oceans (Ng et al., 2008); however, it

is poorly known in the fossil record, and only

Arges de Haan, 1835, Cryptolutea, and Typhlo-

carcinus Stimpson, 1858 are recognized as fos-

sils (de Haan, 1835; Karasawa, 1993; Hu and

Tao, 1996).

Cryptolutea warreni Karasawa and Kato,

new species

(Fig. 4A–D)

Material examined: MGB-CF0015 (holotype),

MGB-CF0016 (paratype), and NMNS PA16394

from DVO-11.

Diagnosis: Moderate-sized Cryptolutea. Cara-

pace rounded rectangular. Front protruded anteri-

orly, downturned, with shallow, narrow median

groove; frontal margin rimmed with 2 rounded

lobes. Upper orbital margin granular, rimmed,

separated from frontal margin by weak notch.

Anterolateral margin strongly convex, granular,

divided into 4 lobes by very shallow notches;

short, oblique groove extending from last two an-

terolateral notches on epibranchial region. Junc-

tion between anterolateral and posterolateral

margins rounded. Posterolateral margin about

equal to anterolateral margin, slightly convex.

Posterior margin nearly straight, granular,

rimmed. Dorsal surface moderately arched trans-

versely, nearly ﬂat longitudinally; surface densely

covered with ﬁne granules and ﬁne granular lines

irregularly arranged. Regions poorly deﬁned.

Branchial regions undifferentiated. Chelipeds un-

equal, dissimilar; right cheliped slightly larger

than left. Carpus ﬁnely tuberculated on lateral

surface. Palm short; lateral surface with rows of

tubercles; dorsal surface ﬁnely tuberculate; ven-

tral margin fringed with scattered tubercles. Both

ﬁngers short.

Etymology: In honor of the late Warren Blow,

a specialist on fossil crabs who was our friend.

Description: Moderate-sized carapace for

Cryptolutea. Carapace rounded rectangular in

outline, length about 75% carapace width, widest

at mid-length. Fronto-orbital margin about 54%

carapace width. Front protruded anteriorly,

64 H. Karasawa et al.

downturned, with shallow, narrow median de-

pression; frontal margin rimmed with 2 rounded

lobes. Upper orbital margin granular, rimmed,

separated from frontal margin by weak notch.

Anterolateral margin strongly convex, granular,

divided into 4 lobes by very shallow notches; 1st

lobe ($outer orbital angle) broadly triangular;

2nd lobe broadest of all lobes; short, oblique

groove extending from last two anterolateral

notches on epibranchial region. Junction between

anterolateral and posterolateral margins rounded.

Posterolateral margin about equal to anterolateral

margin, slightly convex. Posterior margin nearly

straight, granular, rimmed, about 65% carapace

width. Dorsal surface moderately arched trans-

versely, nearly ﬂat longitudinally; surface densely

covered with ﬁne granules and ﬁne granular lines

irregularly arranged, but in small specimen sur-

face smoother. Regions poorly deﬁned. Protogas-

tric regions separated from surrounding regions

by shallow, broad grooves. Mesogastric region

separated from cardiac region by narrow groove

and from branchial regions by rather deep

grooves. Hepatic regions separated from

branchial regions by obtuse groove. Branchial re-

gions undifferentiated. Branchiocardiac groove

shallow. Intestinal region narrow.

Chelipeds unequal, dissimilar; right cheliped

slightly larger than left one. Carpus of right che-

liped poorly preserved; lateral surface ﬁnely tu-

berculate. Palm of right cheliped about 1.25

times longer than high; lateral surface slightly

convex, with 2 rows of tubercles proximally; dor-

sal surface gently convex, ﬁnely tuberculate; ven-

tral margin sinuous, fringed with scattered tuber-

cles. Fixed ﬁnger of right cheliped about half

palm length, with acutely pointed tip; occlusal

margin bearing 3 broadly triangular teeth, middle

one largest; ventral margin gently convex; ven-

tro-lateral margin keeled. Dactylus of right che-

liped slightly longer than palm, gently curved

downward; lateral surface with a row of pits me-

dially; occlusal margin with 3 broad teeth. Palm

of left cheliped about 1.32 times longer than

high; 4 rows of irregular tubercles present on lat-

eral surface, diminishing in length toward dorsal

margin, upper 3 rows running from proximal

margin to median part; dorsal margin gently con-

vex, ﬁnely tuberculate; ventral margin sinuous,

armed with irregular tubercles. Fixed ﬁnger

about 60% palm length, with acutely pointed tip;

occlusal margin with irregular, triangular teeth,

proximal one highest, middle one broadest; ven-

tral margin nearly straight; ventro-lateral margin

keeled, crenulated. Dactylus broken.

Remarks: The present new species is readily

distinguished from Cryptolutea litoralis by hav-

ing the carapace densely covered with ﬁne gran-

ules and irregularly arranged granulated lines. In

C. litoralis the carapace is punctate. The antero-

lateral teeth in C. litoralis are well separated, but

C. warreni has anterolateral lobes divided by

weak notches. Additionally, the carapace regions

of C. litoralis are well deﬁned whereas in C. war-

reni they are poorly deﬁned.

Family Xanthidae MacLeay, 1838, sensu

Karasawa and Schweitzer, 2006

Subfamily Xanthinae MacLeay, 1838

Genus Liagore de Haan, 1833

Type species: Cancer (Liagore) rubromaculata

de Haan, 1835, by monotypy.

Included fossil species: Liagore rubromaculata

(de Haan, 1835) (also extant).

Remarks: Most recently, Ng and Naruse (2007)

described the third extant species, Liagore pul-

chella Ng and Naruse, 2007, from Vanuatu.

Among these, L. rubromaculata is the only

known as fossil species.

Liagore rubromaculata (de Haan, 1835)

(Fig. 5G–L)

Material examined: MGB-CF0017?0020 and

NMNS PA16396 from DVO-11.

Remarks: This species is abundant in sand-

stone exposed at DVO-11. Liagore rubromacula-

ta is also found in the Pliocene–Pleistocene de-

posits of Taiwan (Hu and Tao, 1996).

Neogene and Quaternary Crustaceans from Philippines 65

66 H. Karasawa et al.

Fig. 5. A, B. Eucrate sp., MGB-CF0024, loc. DVO-11, A, ventral; B, dorsal view. C. Carcinoplax purpurea

Rathbun, 1914, MGB-CF0025, loc. DVO-11, carapace, dorsal view. D. Carcinoplax purpurea Rathbun, 1914,

MGB-CF0026, loc. DVO-11, carapace, dorsal view. E, F. Carcinoplax purpurea Rathbun, 1914, MGB-

CF0026, loc. DVO-11, E, dorsal; F, ventral view. G, H. Liagore rubromaculata (de Haan, 1835), MGB-

CF0017, loc. DVO-11, female, G, ventral; H, dorsal view. I, J. Liagore rubromaculata (de Haan, 1835),

MGB-CF0018, loc. DVO-11, male, I, dorsal; J, ventral view. K, L. Liagore rubromaculata (de Haan, 1835),

MGB-CF0019, loc. DVO-11, female, K, dorsal; L, ventral view. M, N. Demania sp., MGB-CF0023, loc.

DVO-11, carapace, M, frontal; N, dorsal view. Scale bars$1cm.

Genus Demania Laurie, 1906

Type species: Demania splendida Laurie,

1906, by original designation.

Included fossil species: Demania caltripes (Al-

cock, 1898) (as D. reynaudi caltripes (Alcock,

1898) in Hu and Tao (1996)) (also extant); D.

chayiensis (Hu, 1981); D. pilipinas Karasawa and

Kato, new species; D. reynaudi (H. Milne Ed-

wards, 1834) (also extant); D. scamberrima

(Walker, 1887) (also extant); D. wardi Garth and

Ng, 1985 (also extant); D. sp. in this paper.

Remarks: Demania chayiensis (Hu, 1981), a

previously known extinct species was recorded

from the upper Miocene–lower Pliocene of Tai-

wan (Hu, 1981; Hu and Tao, 1996). Four extant

species, D. caltripes and D. reynaudi from the

Pliocene–Pleistocene of Taiwan (Hu and Tao,

1996), D. wardi from the Pleistocene of New He-

brides (Garth and Ng, 1985), and D. scaberrima

from the Pliocene of Java (Van Straelen, 1938),

are recognized as fossils.

Demania pilipinas Karasawa and Kato,

new species

(Fig. 4H–K)

Material examined: MGB-CF0021 (holotype),

MGB-CF0022 (paratype), and NMNS PA16397

from DVO-11.

Diagnosis: Carapace pentagonal, length about

80% carapace width, widest at mid-length. Fron-

to-orbital margin about 40% carapace width.

Front strongly protruded anteriorly, bilobed, with

deep V-shaped notch; frontal margin of each lobe

sinuous, outer angle angular. Anterolateral mar-

gin strongly convex without teeth or lobes. Junc-

tion at anterolateral and posterolateral margins

rounded. Posterolateral margin about as long as

anterolateral margin. Posterior margin nearly

straight. Dorsal regions well deﬁned; elevated re-

gions covered with ﬂattened, rounded tubercles.

Carpus and palm of cheliped densely covered

with large, ﬂattened, rounded tubercles on lateral

surface; carpus with large, erect tubercles on dor-

sal margin; palm with 4–5 tuberculate spines on

dorsal margin. Lateral surface of meri of

pereiopods ﬁnely granulate; dorsal and ventral

margins crested, ﬁnely serrated.

Etymology: The speciﬁc name is derived from

the word, “Pilipinas”, meaning Philippines in the

Pilipinese Language.

Description: Carapace pentagonal in outline,

length about 80% carapace width, widest at mid-

length. Fronto-orbital margin about 40% cara-

pace width. Front protruded anteriorly, bilobed,

medially projected anteriorly, with deep V-shaped

axial notch; frontal margin of each lobe sinuous,

outer angle angular, sloping towards inner orbital

angle. Upper orbital margin narrow, concave,

ﬁnely dentate, without orbital ﬁssures. Anterolat-

eral margin strongly convex, without teeth or

lobes. Junction at anterolateral and posterolateral

margins rounded. Posterolateral margin about as

long as anterolateral margin. Posterior margin

nearly straight, slightly shorter than fronto-or-

bital margin. Dorsal surface gently convex longi-

tudinally and transversely; regions well deﬁned,

elevated regions covered with ﬂattened, rounded

tubercles which vary in size; 1F, 2F, and 1M re-

gions divided into two by median deep groove;

1F regions extremely narrow, smooth; 2F region

separated from 1M by shallow groove; 2L- and

3L regions, and 1R and 2R regions united.

Chelipeds symmetrical. Carpus densely cov-

ered with large, ﬂattened, rounded tubercles on

lateral surface; dorsal margin with large, erect tu-

bercles; mesiodistal spine large, ﬁnely tuberculat-

ed. Palm much longer than high; lateral surface

densely covered with large, ﬂattened, rounded tu-

bercles; dorsal margin with 4–5 tuberculate

spines. Fixed ﬁnger about half palm length, orna-

mented with 2 longitudinal rows of rounded tu-

bercles on lateral surface; occlusal margin with 4

blunt teeth. Dactylus about as long as ﬁxed ﬁn-

ger; occlusal margin with 4 blunt teeth; lateral

surface and dorsal margin smooth except for

ﬁnely tuberculate proximal part.

Meri of pereiopods ﬂattened, much longer than

high; lateral surface ﬁnely granulate; dorsal and

ventral margins crested, ﬁnely serrated.

Remarks: The above-mentioned terminology

Neogene and Quaternary Crustaceans from Philippines 67

used the dorsal carapace regions followed Dana

(1852). The present new species resembles the

extant Demania japonica Guinot, 1997, from the

South China Sea, East China Sea, and Japan, but

differs in that the tubercles on the dorsal carapace

surface are rather large, ﬂattened and not spinose,

and the carpi and propodi of the both chelipeds

are ornamented with large, ﬂattened, rounded tu-

bercles. The smooth anterolateral margins of the

carapace readily distinguish this species from the

extinct Demania chayiensis.

Demania sp.

(Fig. 5M, N)

Material examined: MGB-CF0023 from DVO-

11.

Remarks: The specimen is represented by an

internal mould of the carapace; therefore, de-

tailed carapace ornamentations are unknown.

Additionally, the frontal region is broken. The

well developed anterolateral teeth and a wider

carapace readily distinguish this species from

Demania chayiensis and D. pilipinas. Ornamen-

tation of a posterior carapace region of the pre-

sent species resembles that of two extant species,

D. raynaudi and D. scaberrima. More complete

material will be necessary to conﬁrm identiﬁca-

tion of this species.

Superfamily Goneplacoidea MacLeay, 1838,

sensu Karasawa and Schweitzer, 2006

Family Euryplacidae Stimpson, 1871,

sensu Karasawa and Schweitzer, 2006

Genus Eucrate de Haan, 1835

Type species: Cancer (Eucrate) crenatus de

Haan, 1835, by monotypy.

Included fossil species: see Karasawa and Kato

(2003).

Eucrate sp.

(Fig. 5A, B)

Material examined: MGB-CF0024 from DVO-

11.

Remarks: The present specimen is a broken

carapace associated with the thoracic sternum

and male abdomen. The detailed characters of

the fronto-orbital and anterolateral margins are

obscure. The speciﬁc identiﬁcation of this

species awaits the discovery of well-preserved

carapaces.

Family Goneplacidae MacLeay, 1838, sensu

Karasawa and Schweitzer, 2006

Genus Carcinoplax H. Milne Edwards, 1852

Type species: Cancer (Crutonotus) longimanus

de Haan, 1833, by subsequent designation of

Glaessner (1929).

Included fossil species: see Karasawa and Kato

(2003).

Remarks: Karasawa and Kato (2003) made the

list of previously known fossil species of Car-

cinoplax. However, two species are lacking in

their list: Carcinoplax wuhsini Hu and Tao, 1999,

from the Oligocene–Miocene of Taiwan; and

Carcinoplax prepurpurea Hu and Tao, 2000,

from the Pliocene of Taiwan. Most recently, Cas-

tro (2007) redeﬁned Carcinoplax and transferred

several members previously assigned to the

genus to his new genera, Entricoplax, Menoplax,

Pycnoplax, and Thyraplax. The recognition of

these ﬁve genera is mainly based upon characters

of sternal sutures 6/7, eyes, vulva, and gonopod 1

(Castro, 2007). However, it is very difﬁcult to

adapt his classiﬁcation for the extinct species be-

cause these characters are not usually preserved

within the fossils. Therefore, it is considered best

to place the extinct species within Carcinoplax

sensu lato for the time being.

68 H. Karasawa et al.

Carcinoplax purpurea Rathbun, 1914

(Fig. 5C–F)

Material examined: MGB-CF0025-0028,

NMNS PA16398 from DVO-11.

Remarks: This species is also recorded from

the upper Pliocene of Taiwan (Hu and Tao, 1996;

as Carcinoplax linae Hu and Tao, 1996, syn-

onymised with C. purpurea by Karasawa and

Kato (1993)).

Family Hexapodidae Miers, 1886

Genus Hexapus de Haan, 1835

Type species: Cancer sexpes Fabricius, 1798,

by designation in ICZN Opinion 85.

Neogene and Quaternary Crustaceans from Philippines 69

Fig. 6. A, B. Pinnixa sp., MGB-CF0032, carapace, loc. DVO-11, A, dorsal; B, frontal view. C, D. Hexapus

granuliformis Karasawa and Kato, new species, MGB-CF0029 (holotype), loc. DVO-11, C, dorsal; D, frontal

view. E, F. Podophthalmus vigil (Fabricius, 1798), MGB-CF0030, loc. DVO-11, E, dorsal; F, ventral view.

Scale bars$1cm.

Included fossil species: Hexapus anfractus

(Rathbun, 1909) (also extant); H. decapodus

(Morris and Collins, 1991); H. granuliformis

Karasawa and Kato, new species; H. nakajimai

Imaizumi, 1959; H. pinfoldi Collins and Morris,

1978.

Hexapus granuliformis Karasawa and Kato,

new species

(Fig. 6C, D)

Material examined: MGB-CF0029 (holotype)

from DVO-11.

Diagnosis: Moderate-sized Hexapus. Carapace

trapezoidal, much wider than long, length about

60% carapace width. Fronto-orbital margin about

30% carapace width. Posterior margin slightly

convex, about 90% carapace width. Dorsal sur-

face covered with ﬁne granules, moderately con-

vex longitudinally, slightly convex transversely.

Regions poorly deﬁned. Cervical and branchio-

cardiac grooves moderately deﬁned.

Etymology: The trivial name refers to the ﬁne-

ly granulated dorsal surface of the carapace.

Description: Moderate-sized carapace. Cara-

pace trapezoidal in outline, wider than long,

length about 60% carapace width, widest at pos-

terolateral angle. Fronto-orbital margin about

30% carapace width. Front narrow, about 15%

carapace width, weakly protruded anteriorly,

strongly downturned, slightly widening distally,

with weak median depression; frontal margin

straight, rimmed. Orbit small, weakly rimmed.

Anterolateral margin gently convex, granular,

rimmed, not clearly demarcated from posterolat-

eral margin. Posterolateral margin nearly straight,

granular, rimmed, divergent posteriorly. Postero-

lateral re-entrant well developed with low, broad

lobe. Posterior margin slightly convex, about

90% carapace width. Dorsal surface covered with

ﬁne granules, moderately convex longitudinally,

slightly convex transversely. Regions poorly de-

ﬁned. Cervical groove weakly developed anteri-

orly, moderately developed posteriorly. Branchio-

cardiac groove moderately deﬁned. Hepatic re-

gion weakly deﬁned by shallow grooves. Epi-

branchial region slightly vaulted and meso- and

metabranchial regions differentiated. Intestinal

region narrow, ﬂattened.

Remarks: The new species may be similar to

the extant Hexapus sexpes, but differs in having

the ﬁnely granulated dorsal surface. The cervical

and branchiocardiac grooves in the newspecies

are deeper than those in H. sexpes. The ﬁnely

granulated carapace readily distinguishes this

species from two Miocene species, Hexapus de-

capodus (Morris and Collins, 1991) from

Sarawak and Sabah and H. nakajimai Imaizumi,

1959, from Japan. The carapace in H. granuli-

formis is much narrower than that of H. decapo-

dus and H. nakajimai.

Superfamily Portunoidea Raﬁnesque, 1815

Family Portunidae Raﬁnesque, 1815,

sensu Karasawa et al., 2008

Subfamily Podophthalminae Dana, 1851

Genus Podophthalmus Lamarck, 1801

Type species: Podophthalmus [sic] spinosus

Lamarck, 1801 ($Portunus vigil Fabricius,

1798), by monotypy.

Included fossil species: Podophthalmus

fusiformis Morris and Collins, 1991; P. taiwani-

cus Hu and Tao, 1985; P. vigil (Fabricius, 1798)

(also extant).

Podophthalmus vigil (Fabricius, 1798)

(Fig. 6E, F)

Material examined: MGB-CF0030, MGB-

CF0031, NMNS PA16399 from DVO-11.

Remarks: The fossil occurrence of this species

has previously been reported from the Pliocene

of Java (Martin, 1883), the Pleistocene of

Sarawak (Collins et al., 2003), the Pleistocene of

Guam (Kesling, 1958; Schweitzer et al., 2002),

the Pleistocene of Taiwan (Hu and Tao, 1996),

the Pleistocene of Japan (Karasawa et al., 1995),

and the Holocene of Australia (Etheridge and

McCulloch, 1916).

70 H. Karasawa et al.

Superfamily Pinnotheroidea de Haan, 1833

Family Pinnotheridae de Haan, 1833

Subfamily Pinnotherelinae Alcock, 1990

Genus Pinnixa White, 1846

Type species: Pinnotheres cylindricum Say,

1818, by monotypy (ICZN Opinion 85).

Included fossil species: see Feldmann et al.

(2005).

Remarks: Feldmann et al. (2005) recognized

six species of Pinnixa in the fossil record and ex-

cluded three fossil species, Pinnixa aequipuncta-

ta Morris and Collins, 1991, P. omega Morris

and Collins, 1991, and P. microgranulosa Collins

et al., 2003, from the Neogene of Brunei, Sabah

and Sarawak. They suggested that these three

species might be referred to Tetrias Rathbun,

1898. However, the paratype specimen (BM IC

274) of P. microgranulosa ﬁgured by Collins et

al. (2003, pl. 7, ﬁg. 6) has great similarities with

an immature female specimen (Morris and

Collins, 1991, ﬁg. 57) of Orthakrolophos bittneri

(Morris and Collins, 1991) (see Schweitzer and

Feldmann, 2001, p. 335). The generic status of

only Pinnixa omega is retained in that the cara-

pace is much wider than long and the dorsal sur-

face is not punctuated.

Pinnixa sp.

(Fig. 6A, B)

Material examined: MGB-CF0032 from DVO-

11.

Remarks: A poorly preserved carapace repre-

sents the present species. This species resembles

Pinnixa omega, but differs in that the large-sized

carapace is more ﬂattened with strongly inﬂated

anterolateral margins. More complete material

will be necessary to conﬁrm identiﬁcation of this

species. The carapace of this species may be sim-

ilar to that of the hexapodid Hexapus granuli-

formis. However, the more ﬂattened carapace, the

well deﬁned gastric regions, and the strongly in-

ﬂated anterolateral margins are readily distin-

guished this species from H. granuliformis.

Acknowledgments

We thank C. E. Schweitzer (Kent State Univer-

sity, U.S.A.) for reading carefully the systematic

part of this paper and T. Naruse (National Uni-

versity of Singapore) for providing useful com-

ments on some fossil crabs. We also thank E.

Mula (MGB), R. Ancog (MGB), W. Mago

(MGB), E. Azurin (MGB), C. Neis (Fortune Ce-

ment Co. Ltd.) and R. Guinto (Fortune Cement

Co. Ltd.) for their assistance in ﬁeld work, S.

Laserna who introduced us to the fossil locality

in Batangas, and H. Komatsu (NMNS) for review

of the manuscript. H. C. Ramos, the director of

MGB, is also acknowledged for permitting us to

conduct this research. This study was ﬁnancially

supported by the National Museum of Nature

and Science and by a Grant-In-Aid for the Scien-

tiﬁc Research from the Japan Society for the Pro-

motion of Science (no. 18253007).

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A new species of Galene de Haan, 1833 (Galenidae: Brachyura) from the middle Miocene of Zagros Mountains, Iran

Article

Full-text available

Apr 2022

A new species of Galene de Haan, 1833 (Decapoda: Brachyura: Galenidae) is described from the Mid Miocene (Langhian) strata of the Mishan Formation exposed at two localities in Hormozgan Province, Zagros Mountains, Iran. The galenid species, Galene dashtbani n. sp. is considered as the oldest representative of the genus and simultaneously the westernmost occurrence of all fossil and extant congeners known to date. The genus is currently limited to the Indo-West Pacific region. The present fossil occurrence of the genus suggests a Tethyan origin and subsequent migration eastward.

On some Paleogene and Neogene crabs of Kachchh, Western India.

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Jan 2013

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Sep 2016
Geodiversitas

A new set of Paleocene and Eocene decapod crustaceans is described from the Kirthar Range of Pakistan. Two new ghost shrimps (Crustacea, Decapoda, Callianassidae) are described: Neocallichirus khadroensis Hyžný & Charbonnier, n. sp. from the Paleocene (Danian, Khadro Formation) of Gawar Band, Ranikot District, and Neocallichirus lakhraensis Hyžný & Charbonnier, n. sp. from the Early Eocene (Ypresian, Lakhra Formation) of Rbod Nala, Jhirak District. Both new species exhibit chelipeds which are morphologically surprisingly close to extant Neocallichirus karumba (Poore & Griffin, 1979) from the Indo-West Pacific. A group of species sharing this same cheliped morphology is provisionally called the “karumba group” based on Neocallichirus karumba, best documented species. The “karumba group” encompasses seven fossil species: the two new Pakistani species, Neocallichirus tuberculatus (Lőrenthey in Lőrenthey & Beurlen, 1929) n. comb. from the Eocene of Hungary, Neocallichirus borensis Beschin, De Angeli, Checchi & Mietto, 2006 from the Eocene of Italy, Neocallichirus birmanicus (Noetling, 1901) n. comb. from the Miocene of Myanmar, Neocallichirus dijki (Martin, 1883) from the Miocene of Java and Philippines, and the subfossil Neocallichirus maximus (A. Milne-Edwards, 1870) from Thailand. Based upon the extant and fossil occurrences, it is difficult to reconstruct migration pattern of the “karumba group”. For now, it can be concluded, that at the genus level, a relative homogeneity of the ghost shrimps is observed between the Eastern and the Western Tethyan regions, as already suggested by Merle et al. (2014) for the assemblage of volutid gastropods from the Lakhra Formation.

Holocene Crab Consumption at Watinglo, Papua New Guinea

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Dec 2023

Hendri A. F. Kaharudin

Malacostraca are commonly found in coastal archaeological sites in the Indo-Pacific; both swimming and terrestrial crabs. This archaeological evidence as well as ethnographic studies indicate that they formed an important component of the human diet in the past as they still do in many regions today. However, they often go unidentified and largely unanalysed in archaeological research due to the highly fragmentary state of the remains and the associated difficulties posed for identification and quantification. Here we discuss these issues and provide an example of crab remains recovered from a coastal archaeological site in Papua New Guinea, Watinglo shelter, with an occupation record spanning from the Terminal Pleistocene to historic times. We also briefly compare these finding to crab remains from archaeological sites of similar antiquity in Indonesia and the Philippines.

Crustáceos braquiuros del Mioceno de las secciones Konar-Takteh y Ahram, Irán suroccidental

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Aug 2013

Five species of brachyuran Crustacea from the Middle Miocene are described from the Guri Member of the Mishan Formation in southwestern Iran. The species include the leucosioids Leucosia persica; Seulosia sp. cf. S. rhomboidalis; Myra sp.; the portunoid Portunus withersi and the xanthid Demania ahramensis n. sp. The paleobiogeographic affinities of these genera reinforce previous interpretations of a wide distribution between the Persian Gulf and the Indopacific regions.

A new species of ghost shrimp (Decapoda, Thalassinidea, Callianassidae) from the Late Cretaceous (Cenomanian) of Agadir (W Morocco)

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Jan 2011

We report Neocallichirus agadirensis n. sp. (Decapoda, Thalassinidea, Callianassidae) from the Late Cretaceous (Cenomanian) of Agadir (W Morocco). This species has been described based upon four specimens that preserve ischium, merus, carpus, propodus, and dactylus of the great cheliped. The new species represents the first report of Neocallichirus in Africa and the oldest species known to date belonging to this genus.

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The oldest freshwater crabs: claws on dinosaur bones

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Dec 2019

With approximately 1,500 extant species, freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans. Nevertheless, their fossil record is extremely limited: only Potamidae, Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far. This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France), in close association with large vertebrate remains. In addition to (1) the systematic assignment of these claws, the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils, and (3) the likely scenario for their auto/allochthony in the Velaux fluvial system. These claws belong to a new taxon, Dinocarcinus velauciensis n. gen. n. sp., referred to as Portunoidea sensu lato, a group of “true” crabs nowadays linked to marine systems. However, the faunal assemblage, the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology, making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma. Either as primary or as secondary freshwater crabs, the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time, as early as the Campanian.

The oldest freshwater crabs: claws on dinosaur bones

Preprint

Full-text available

Aug 2019

With approximately 1,500 extant species, freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans. Nevertheless, their fossil record is extremely limited: only Potamidae, Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far. This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France), in close association with large vertebrate remains. In addition to (1) the systematic assignment of these claws, the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils, and (3) the likely scenario for their auto/allochtony in the Velaux fluvial system. These claws belong to a new taxon, Dinocarcinus velauciensis n. gen. n. sp., referred to as Portunoidea sensu lato, a group of "true" crabs nowadays linked to marine systems. However, the faunal assemblage, the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology, making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma. Either as primary or as secondary freshwater crabs, the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time, as early as the Campanian.

The identify of Cancer (Arges) parallelus De Haan, 1833 (Decapoda: Brachyura: Pilumnidae), a fossil crab described from Japan during the 19th century

Article

Jul 2019
J CRUSTACEAN BIOL

The revised diagnosis and description are given for Cancer (Arges) parallelus De Haan, 1833, a fossil crab first described from Japan during the 19th century. The neotype of this species from the Holocene (about 9,000-5,000 ybp) Nanyo Formation of Ise Bay, central Japan, is herein designated. The monotypic genus Arges De Haan, 1833 belongs to the pilumnid sub-family Rhizopinae Stimpson, 1858 and has close affinities with Typhlocarcinus Stimpson, 1858, a genus of Rhizopinae.

Systema Brachyurorum: Part I. An annotated checklist of extant Brachyuran crabs of the world

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Jan 2008
RAFFLES B ZOOL

A new genus and five new species of Callianassidae (Crustacea: Decapoda: Thalassinidea) from Northern Australia

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Dec 1988

Katsushi Sakai

Marine Decapod Crustacea of Southern Australia: A Guide to Identification

Book

Jan 2004

Gary C. B. Poore

This book is a comprehensive guide to the identification of 800 species of decapod and stomatopod crustaceans from southern Australian marine waters. It is liberally illustrated with more than 1000 line drawings giving good views of many species as well as diagnostic illustrations. Details for each species include the authority, year of description, sometimes a common name, diagnosis, size, geographical distribution, and ecological and depth distribution. The chapter on the Stomatopoda is by Shane Ahyong. Sections within each chapter are hierarchical, species within genera, within families (often with subfamilies as well). Identification is achieved through the use of dichotomous keys adapted from many originally published in the primary literature, or developed from scratch. Some keys are to all Australian taxa but most are to southern Australian taxa only. The information in this book derives from over 200 years of collecting in southern Australian environments, from the intertidal to the deep sea, and publications in numerous journals in several languages. More than 800 of these papers and books are cited. Winner of the 2005 Whitley Award for Systematics.

Nomenclator entomologicus secundum Entomologiam Systematicum ill. Fabricii adjectis speciebus recens detectis et varietatibus

Article

F. Weber

Revision of the American Callianassidae (Crustacea, Decapoda, Thalassinidea)

Article

Jan 1991

Gx136, ISI:A1991GX13600011

Neogene crabs from Brunei, Sabah and Sarawak

Article

Jan 1991

Thirty-six species of fossil crab are described and figures from the Neogene of Sabah, Sarawak and Brunei. The following 3 genera, 31 species and 3 subspecies are new: Dorippe frascone tuberculata, Calappa sexaspinosa, Podophthalmus fusiformis, Charybdis feriata bruneiensis, Portunus obvallatus, P. woodwardi, Galene stipata, Parthenope (Rhinolambrus) sublitoralis, Ampliura (gen. nov.) simplex, Drachiella guinotae, Iphiculus granulatus, I. miriensis, I. sexspinosus, Leucosia longiangulata, L. serenei, L. tutongensis, Myra brevisulcata, M. subcarinata, M. trispinosa, Nucia borneoensis, N. calculoides, N. coxi, Nucilobus (gen. nov.) symmetricus, Pariphiculus gselli beetsi, P. papillosus, P. verrucosus, Philyra granulosa, Typilobus marginatus, Palaeograpsus bittneri, Pinnixa aequipunctata, P. omega, Prepaeduma (gen. nov.) decapoda, Xenophthalmus subitus, Macrophthalmus (Maerotis) wilfordi. -Authors

Observations sur les affinités zoologiques et la classification naturelle des Crustacés

Article

H. Milne Edwards

A new genus and species of callianassid ghost shrimp from Kyushu, Japan (Decapoda : Thalassinidea)

Article

Jun 2000

Gary C. B. Poore

Grynaminna tamakii, new genus, new species, is described from specimens taken in a small area near the south end of Shimabara Peninsula, western Kyushu, Japan. The new genus shares with the eight known callichirine callianassid genera presence of a dorsal oval, maxilliped 3 with ovate propodus and slender dactylus, and pleopods 3-5 with "stubby" appendices internae. It most resembles Callichirus but differs in possessing an elongate-oval uropodal endopod, absence of a meral hook an the major cheliped, and absence of dorsal sculpture on abdominal somites 3-5.

Diagnoses préliminaires de 34 espèces et variétés nouvelles, et de 2 genres nouveaux de Décapodes de la Mer Rouge

Article

Jan 1905

G. Nobili

Materials for a carcinological fauna of India. No. 3. The Brachyura Cyclometopa. Part I. The family Xanthidae

Article

A. Alcock

Neogene and Quaternary ghost shrimps and crabs (Crustacea: Decapoda) from the Philippines

Abstract and Figures

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