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Original Article • Nauplius 34 • 2026 • https://doi.org/10.1590/2358-2936e20260605 linkcopy

Open-access Extant and fossil species of Karumballichirus (Decapoda: Axiidea: Callichiridae)

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

Abstract

Two extant species of Indo-West Pacific ghost shrimps of the genus Karumballichirus Poore, Dworschak, Robles, Mantelatto and Felder, 2019 that had been previously synonymized, are distinguished on the basis of molecular and morphological criteria. Karumballichirus karumba (Poore and Griffin, 1979) from northern Australia, Papua New Guinea, Singapore, and Thailand differs from Karumballichirus kempi (Sakai, 1999) from India, Indonesia and Taiwan in the shape of the adult cheliped, telson, and uropods, and in reaching little over half the length of K. kempi. Ten fossil species of Karumballichirus, six in new combinations, dating from the Paleocene (Danian) to Holocene are listed: Karumballichirus birmanica (Noetling, 1901) n. comb., Miocene; Karumballichirus dijki (Martin, 1883) n. comb., Miocene; Karumballichirus lakhraensis (Hyžný and Charbonnier in Hyžný, Charbonnier, Merle, Lashari, Bartolini and Métais, 2016), early Eocene (Ypresian); Karumballichirus khadroensis (Hyžný and Charbonnier in Hyžný, Charbonnier, Merle, Lashari, Bartolini and Métais, 2016), Paleocene (Danian); Karumballichirus pseudoniloticus (Lőrenthey in Lőrenthey and Beurlen, 1929) n. comb., middle Eocene; Karumballichirus pustulatus (Withers, 1926) n. comb., ?Eocene-Oligocene; Karumballichirus trechmanni (Withers, 1924) n. comb., Eocene; Karumballichirus tuberculatus (Lőrenthey in Lőrenthey and Beurlen, 1929), middle Eocene; Karumballichirus vidali (Vía Boada, 1959) n. comb., early Eocene, including the subfossil Karumballichirus maximus (A. Milne-Edwards, 1870) which is not a senior synonym of any modern species.

Keywords
Fossils; ghost shrimps; Indo-West Pacific; taxonomy

INTRODUCTION

Dworschak (2008) reviewed the taxonomic history of three species of similar Indo-West Pacific callichirid ghost shrimps, Callianassa maxima A. Milne-Edwards, 1870, Callianassa karumbaPoore and Griffin, 1979, and Neocallichirus kempiSakai, 1999, placing all in Neocallichirus Sakai, 1988. The first species is based on a chela recovered from a canal in Thailand. The description of C. karumba was based on a single male from tropical Australia. Sakai (1999) selected a post-ovigerous female from Bangka, Indonesia, as the holotype of his new species N. kempi, while appearing to intend a new name for the extant Indian species that Kemp (1915) had identified as C. maxima. Sakai listed but did not designate as paratypes, two specimens from India. Further Indian extant examples of “C. maxima” from India have been figured by Pillai (1954) and Daniel (1981). Dworschak (2008) believed C. karumba and N. kempi were synonymous and extensively redescribed the species as N. karumba (Poore and Griffin, 1979) on the basis of numerous specimens from the West Pacific.

Dworschak’s (2008) synonymy was disputed by Sakai (2011). A fourth species, similar to these, Neocallichirus thalesapensis Sakai and Lheknim, 2014 was described later. The similarities between these extant species and older fossils were realized by Hyžný et al. (2016) who treated them as the “Karumba group” of Neocallichirus.

Poore et al. (2019) introduced a new genus name, Karumballichirus Poore, Dworschak, Robles, Mantelatto and Felder, 2019, for this group justifying it on a revisionary study of callianassoid phylogeny (Robles et al., 2020). This study analysed molecular sequences (18S, 16S, 12S, H3 genes) from six individuals identified by the authors as Karumballichirus karumba (Poore and Griffin, 1979), finding that they fell into two sister clades, one from Singapore and another from Taiwan, with significant genetic separation, suggesting the probability of more than one extant species in the southwestern Pacific.

This contribution reappraises the extant species of Karumballichirus in the light of genetic and morphological evidence. Taxonomy of the “Karumba group” of fossil species is also reviewed.

MATERIAL AND METHODS

We have reassessed (but not listed again) the extensive material listed by Dworschak (2008) from Naturhistorisches Museum, Vienna (NHMW), the Muséum nationale d’Histoire naturelle, Paris (MNHN), National Taiwan Ocean University, Taiwan (NTOU), Zoological Reference Collection, Lee Kong Chian Natural History Museum, National University of Singapore (ZRC); and Queensland Museum, Brisbane, Australia (QM), and by Hyžný et al. (2025). We have also re-examined specimens from Museums Victoria, Melbourne, Australia (NMV), the Western Australian Museum, Perth (WAM), and the Natural History Museum, London (NHMUK). Tissue samples are from the collection of the Department of Biology, University of Louisiana Lafayette, USA (ULLZ) derived in turn from museum specimens from NHMW, NTOU, and ZRC. They were processed as explained by Robles et al. (2020); GenBank numbers can be found in Supplementary Table S1 in Robles et al. (2020).

All figures of extant species are drawn to the same scale from camera lucida pencil drawings converted to vector illustrations in Adobe Illustrator®. Dimensions are given as carapace length (cl.) and total length (tl.).

The studied fossil material included specimens listed in Hyžný et al. (2016; 2025) and Hyžný and Zorn (2020). Newly examined material included specimens deposited in Museo Geológico del Seminario de Barcelona, Spain (MGSB) and the Natural History Museum, London; details are given in figure captions. The methods used to examine fossils are essentially those used by Hyžný and Klompmaker (2015). Specimens deposited in MGSB (Figs. 5 and 8) were coated with the ammonium chloride sublimate prior to photography; Nikon Coolpix 7800 was used to document the specimens. Specimens deposited in NHMUK (Figs. 6 and 7) were assessed using Stereo Zoom Microscope ZEISS Axio Zoom.V16 with AxioCam HRC and a Canon EOS 750D camera with EF 50 mm f/2.5 Compact Macro Lens.

SYSTEMATICS

Callichiridae Manning and Felder, 1991

Karumballichirus Poore, Dworschak, Robles, Mantelatto and Felder, 2019

KarumballichirusPoore et al., 2019: 111.- Robles et al., 2020: supplementary tables 1, 2.

Type species. Callianassa karumbaPoore and Griffin, 1979, by original designation.

Remarks. The genus was erected as part of a molecular and morphological review of Callianassidae and related families. The genus is distinguished from the more species-rich Neocallichirus (see Poore et al., 2019: Robles et al., 2020) by possession of a short maxillipedal 3 exopod, a feature shared with Lepidophthalmus Holmes, 1904. Specimens of Karumballichirus can be readily recognized by the quadrant-shaped tooth at the base of the fixed finger of the major cheliped.

The two sister clades revealed by genetic analysis, within what was initially identified as K. karumba by Robles et al. (2020: fig. 4), were further investigated. Estimates of evolutionary divergence between the sequences of four individuals from Singapore and the two individuals from Taiwan, are for the present limited to 16S and 12S mt sequence data. These genes have proved effective in species differentiation in different taxa including vertebrates like frogs (Dawood et al., 2001; Lemmon et al., 2007) and pit vipers (Malhotra et al., 2011), as well as invertebrates like trematodes (Chan et al., 2022), isopods (Hurtado et al., 2016), crayfish (Munasinghe et al., 2004; Buhay et al., 2007), swimming crabs (Robles et al., 2007), and freshwater crabs (Daniels, 2017).

Genetic divergence within the two populations, as measured by the percentage of number of base differences, was zero or negligible (0-0.25%) for both genes (Tab. 1). Genetic divergence between the two populations from Singapore and Taiwan was significant, 9.26% for the 16S gene, and 6.42-6.67% for the 12S gene (Tab. 1). This level of sequence divergence is typically found among other pairs of congeneric callichirid species. For example, species of GlypturusStimpson, 1866 show sequence divergence from 8.9% to 13.39% for the 12S gene and 7.08-8.44% for 16S (Robles et al. 2020). Sato et al. (2024) showed genetic divergence for the 16S gene between five species of Glypturus of between 6.67% and 17.68%. Species of Callichirus showed sequence divergence from 7.17% to 14.94% for the 12S gene and 5.34-11.39% for 16S (Robles et al., 2020), while species of Lepidophthalmus Holmes, 1904 showed sequence divergence from 5.3% to 10.3% for the 16S gene and 10.47-17.48% for 12S (Felder and Robles, 2015).

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Table 1.
Estimates of evolutionary divergence between sequences from six individuals of Karumballichirus species from Singapore and Taiwan. In the upper half of the table the number of base differences per site between sequences are shown, based on five nucleotide sequences of the 12S mt gene, while in the lower half of the table the number of base differences per site between sequences are shown, based on six nucleotide sequences of the 16S mt gene. All ambiguous positions were removed for each sequence pair (pairwise deletion option). Evolutionary analyses were conducted in MEGA X (Kumar et al., 2018).

We conclude that these genetic differences support the recognition of two extant species in the southwestern Pacific and prompted us to reassess the collections examined by Dworschak (2008) plus some new material. We were able to find consistent morphological differences between populations from Taiwan and Singapore for a few of the characters reviewed, a difficulty confounded by allometric and sexual differences. Dworschak’s synonymy of K. kempi with K. karumba was disputed by Sakai (2011) who noted differences in the male pleopods 1 and 2, and the telson. He remarked that the male pleopod 1 of the holotype of C. karumba is distally W-shaped (Dworschak, 2008: fig. 1i; Poore and Griffin, 1979: fig. 31g). We found the same form of male pleopod 1 in another small (10.9 mm) male from the Northern Territory but a larger male (17.4 mm) (Dworschak, 2008: fig. 3h) showed the fully adult form with two sharp curved apices. Sakai (2011) observed that the appendix interna on the male pleopod 2 is “proximally-fused” in K. kempi and “small” in K. karumba. The appendix interna is terminal on the 8.6 mm holotype (Dworschak, 2008: fig. 1j; Poore and Griffin, 1979: fig. 31h) but lateral to the appendix interna in larger individuals such as a 17.4 mm male from Singapore (Dworschak, 2008: fig. 3i) and in all other males figured by him. These differences between male pleopods are age-dependent and do not distinguish the species.

Sakai (2011) noted that the telson of K. karumba is “trapezoid” while that of K. kempi is “suboval” basing his comparison on the telson of the holotypes, with carapace lengths 8.6 mm and 30 mm respectively. This is largely but subtly true across all ages, the telson of K. karumba being widest near the anterior margin and tapering at least over the posterior half while that of K. kempi is widest at the midpoint. Further subtle differences distinguish the two species but both are highly variable. The uropodal endopod becomes more elongate and narrower with increasing size in both species. The posterior margin is straighter (except in the smallest individuals) in K. kempi than in K. karumba and the slight flexure near the end of the anterior margin in K. kempi is not apparent in K. karumba (cf. Fig. 1D-E, J with Fig. 3). The major cheliped becomes proportionally larger with increasing size. At about cl. 21 mm a semicircular notch appears under the tooth above the fixed finger in the male of K. kempi (Fig. 2A-D) but in the male of K. karumba the notch is a shallow angle (Fig. 1A-C). The notch is smaller and more angled in females of both species (Figs. 1G, H, 2F-I).The dactylar cutting edge of the major cheliped of the largest adults of both sexes of K. kempi is armed with a series of teeth: a proximal group of 1-3 (sometimes fused), a deep hiatus, a rounded tooth near the midpoint, a subdistal tooth or series of teeth, and a recurved apex in the largest individuals (Fig. 2). This edge in K. karumba is less complex: proximal teeth are absent, a shallow hiatus may be present in larger individuals, and the distal half is concave and finely denticulate (Fig. 1A-C, G, H).

We were able therefore to differentiate K. kempi (from Taiwan, Indonesia, and India) from K. karumba (from Australia, Papua New Guinea, and Singapore) on the basis of shapes of the telson, uropod, and major cheliped as detailed above. The maximum size of the two species differs significantly. The holotype of K. kempi has a carapace length of 30 mm (tl. 131 mm); several specimens from Taiwan are in the range 20-24 mm (76-100 mm) while the smallest is 10 mm (37 mm) (Dworschak, 2008). The maximum carapace length of K. karumba is an ovigerous female of 18.6 mm (73 mm), another female is 22 mm (90 mm), and the smallest known is 9 mm (30 mm).

Extant species

Karumballichirus karumba ( Poore and Griffin, 1979 )

(Fig. 1)

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Figure 1.
Karumballichirus karumba (Poore and Griffin, 1979), major chelipeds (lateral faces), tailfans (males on left, females on right). A, D, male, 17.4 mm, Singapore (ZRC 2002.0275); B, male, 17.8 mm, Papua New Guinea (QM W10245); C, F, male, cl. 10.9 mm, Northern Territory, Australia (NTMAG Cr.001289); E, male, 14.5 mm, Papua New Guinea (QM W10240); G, female, 22 mm, Singapore (ZRC 2002.0274); H, female, 7.2 mm, Western Australia (WAM C42970); I, J, holotype ovigerous female, 8.6 mm, Qld, Australia (AM P.24678). Scale bar = 10 mm. C, F, H original; others traced from Dworschak (2008).

Callianassa karumbaPoore and Griffin, 1979: 266, figs. 30, 31.

Glypturus karumba.- Sakai, 1988: 61.- Tudge et al., 2000: 144.- Davie, 2002: 460.

Neocallichirus karumba.- Sakai, 1999: 101.- Sakai, 2005: 180.- Dworschak, 2008: 75-84 (part), figs. 1-3, 6.- Sakai, 2011: 459.

Neocallichirus thalesapensisSakai and Lheknim, 2014: 93-96, figs. 1, 2. (N. syn.)

Karumballichirus karumba.- Poore et al., 2019: 144.- Robles et al., 2020: fig. 4 (part from Singapore).

Karumballichirus thalesapensis.- Poore et al., 2019: 144.

New material examined. Australia, Western Australia, Ashburton River delta, 21.69°S 114.91°E, WAM C42470 (female, cl. 7.2 mm; 2 females, 4.1, 4.4 mm). - Northern Territory, NT Fisheries stn D.166, NTMAG Cr.001289 (male, cl. 10.9 mm). Singapore, St John Island, north lagoon, 1°13.116’N 103°51.079’E, stn SW117, ZRC 2018.0526 (female, cl. 7.6 mm); same locality, stn YB188, ZRC 2018.0525 (female, cl. 9.6 mm).

Diagnosis. Telson widest near anterior margin, tapering at least over posterior half. Uropodal endopod posterior margin convex, anterior margin evenly convex. Major cheliped palm mostly smooth, tuberculate distolaterally in larger individuals, with simple triangular notch under tooth above fixed finger at all sizes; fixed finger rarely with tuberculate ridge on lateral face, without lateral teeth near its base; dactylus shorter than upper margin of palm, cutting edge of adults of both sexes without proximal teeth, sometimes shallow hiatus in larger individuals, distal half concave and finely denticulate. Maximum cl. 22 mm.

Distribution. Northern Australia, Papua New Guinea, Singapore, Thailand (estuarine environments).

Remarks. The holotype is a 34 mm (cl. 8.6) male from northern Queensland, Australia. Dworschak (2008) redescribed this specimen and illustrated other individuals from Singapore and Papua New Guinea - specimens from Taiwan, that he believed belonged to the same species, are here treated as K. kempi. Neocallichirus thalesapensisSakai and Lheknim, 2014 was described from the Songkhla lagoon system, southeastern coast of Thailand, presumably a brackish environment. Type material includes males and females with carapace lengths ranging from 15.5 to 18.6 mm (tl. 66-75 mm). The major cheliped, telson and uropodal endopod are within the range of variability of K. karumba and the species is here synonymized.

Karumballichirus kempi ( Sakai, 1999 )

(Figs. 2, 3)

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Figure 2.
Karumballichirus kempi (Sakai, 1999), major chelipeds (A-E, males, F-I, females). A, B, male, 23.4 mm, India (NHMUK 1938.6.23.3-4), mesial and lateral; C, male, 22.4 mm, Taiwan (NHMW 21937), lateral; D, male, 21.4 mm, Taiwan (NHMW 21936), lateral; E, male, 14.0 mm, Taiwan (NMV J58870), lateral; F, holotype female, 30 mm (ZMB 3353), lateral; G, H, female, 17.3 mm, India (NHMUK 1938.6.23.3-4), mesial and lateral; I, female, 22.3 mm, Taiwan (NTOU 00089), lateral; Scale bar = 10 mm. A, B, E, G, H original; F traced from Sakai (1999); C, D, I traced from Dworschak (2008).

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Figure 3.
Karumballichirus kempi (Sakai, 1999), tailfans (A-C, males, D-E, females). A, male, 24.0 mm, Taiwan (NMV J55324); B, male, 23.4 mm, India (NHMUK 1938.6.23.3-4); C, male, 14.0 mm, Taiwan (NMV J58870); D, holotype female, 30 mm, Indonesia (ZMB 3353; reproduced from Sakai, 1999); E, female, 25.0 mm, Taiwan (NMV J55324); F, female, 17.3 mm, India (NHMUK 1938.6.23.3-4). Scale bar = 10 mm.

Callianassa (Callichirus) maxima.- Kemp, 1915: 252-257, fig. 21, pl. 13 figs. 1-5.- De Man, 1928: 29, 92, 112 (part).- Daniel, 1981: 193-204, pl. 6 figs. 1, 2.

Callianassa maxima.- Pillai, 1954: 23-26, figs. 1-5.

Neocallichirus kempiSakai, 1999: 101-103, fig. 24.- Sakai, 2005: 180.- Sakai, 2011: 459-460.

Neocallichirus karumba.- Dworschak, 2008: 75-84 (part), figs. 4, 5.

Karumballichirus karumba.- Poore et al., 2019: 144.- Robles et al., 2020: fig. 4 (part from Taiwan).- Padate et al., 2022: 198 (list).

Not Callianassa maxima A. Milne-Edwards, 1870: 97-98, pl. 2 fig. 5.

New material examined. Taiwan, Changhua County, Shengang, tidal flat, 24°10’02.52”N 120°27’24.33”E, NMV J55324 (ex NTOU 00089) (male, cl. 24 mm), NMV J58870 (ex MNHN Th1512) (male, cl. 13 mm; female, cl. 24 mm; chela from larger male). NHMW 25376 (female, cl. 28.5 mm), NHMW 25377 (female, cl. 26.7 mm). India, N. Parur, Travancore (now North Paravur, Kerala) NHMUK 1938.6.23.3-4 (male, cl. 23.4 mm; female, cl. 17.3 mm).

Diagnosis. Telson widest at midpoint, suboval. Uropodal endopod posterior margin almost straight, with slight flexure near end of anterior margin. Major cheliped palm tuberculate mesially and laterally in larger individuals, with semicircular notch (about fifth of palm width) between quadrant-shaped tooth and fixed finger in males larger than about cl. 21 mm and larger females; larger males often with curved row of tubercles on distolateral face extending on to fixed finger; dactylus cutting edge of adults armed with series of teeth: a proximal group of 1-3 (sometimes fused), a deep hiatus, a rounded tooth near the midpoint, a subdistal tooth or series of teeth, and a recurved apex in the largest individuals (obsolete in smaller females). Maximum cl. 30 mm.

Distribution. India, Indonesia, Taiwan (estuarine environments).

Remarks. Sakai (1999) illustrated the female holotype of cl. 30 mm from Indonesia and listed two other specimens from N. Parur, Travancore (now North Paravur, Kerala), India. These two are illustrated here for the first time. Shrimps from southern India identified as Callianassa maxima have been thoroughly and consistently described and illustrated. Kemp (1915) illustrated a specimen from Madras (Chennai) and recorded others from Chilka Lake, Pillai (1954) illustrated material from Kayamkulam Lake, Central Travancore (Kerala), and Daniel (1981) figured adults and larvae from Tamil Nadu and Andhra Pradesh where the species is a pest in salt-pans. None of these authors noted the sex of their specimens which might explain some disparity between their illustrations, but all are of adults. None of this material was examined as part of this study.

Fossil species

The fossil record of Karumballichirus is uneven, mainly due to the delicate nature of the majority of cuticular surfaces of the ghost shrimps. Usually, only isolated major cheliped elements are preserved as these are heavily calcified. Occasionally, the cheliped elements are articulated to the extent that the entire cheliped is preserved (i.e., the cheliped disassociation unit). Thus, the attribution of the fossil material to the genus Karumballichirus is based on the evaluation of the characters present on major cheliped only. In Karumballichirus, these are, however, developed in a unique way allowing identification at the genus level even in cases when only fragmentary and/or isolated elements are at hand. The unique set of characters include the subquadrate palm, often tuberculated, with a distinct quadrant-shaped tooth at the base of the fixed finger, dactylus with a dentition consisting of peg-shaped teeth and hooked tip, and merus, longer than high, with the lower margin armed with prominent spines and large proximal hook, often bifid.

The oldest fossil occurrence of the genus is Karumballichirus sp. from the Maastrichtian of Madagascar (Hyžný et al., 2025). Further occurrences are known from Paleocene (K. khadroensis), Eocene (K. lakhraensis, K. pseudoniloticus, K. trechmanni, K. tuberculatus, K. vidali), Oligocene (K. pustulatus), Miocene (K. birmanicus, K. dijki) and subfossil strata (K. maximus).

Karumballichirus birmanicus ( Noetling, 1901 ) n. comb.

Callianassa birmanicaNoetling, 1901: 368, pl. 24 figs 3-5.-Schweitzer et al., 2010: 34 (list).

Remarks. Callianassa birmanica has been reported from the Miocene of Myanmar, based on several isolated major cheliped propodi (Noetling, 1901). Noetling (1901) remarked on similarities between C. birmanica and both C. dijki Martin, 1883 and C. maxima, all species treated here as belonging to Karumballichirus (see below). Noetling (1901) noted that Callianassa birmanica differs from C. dijki in tuberculation on the propodus (coarser in C. birmanica) and arrangement of setal pores on the same element. The taxonomic value of these characters is questionable; nevertheless, without personal examination of the type material of both species, C. birmanica and C. dijki, it is difficult to make any further implications on their possible synonymizing. For the time being, both taxa are treated as separate species within Karumballichirus.

Karumballichirus dijki (Martin, 1883) n. comb.

Callianassa Dijki Martin, 1883: 36, pl. 3 figs 31-33.-Martin, 1895: 57. -Smith, 1913: 293, pl. 19 figs. 3, 4.-Böhm, 1922: 522, pl. 43 figs. 5, 6.

Callianassa frangensBöhm, 1922: 523, pl. 43 fig. 4. (N. syn.)

Callianassa Djiki [sic].-Glaessner, 1929: 79.

Neocallichirus dijki.-Karasawa et al., 2008: fig. 3B, C. -Schweitzer et al., 2010: 38 (list).

Remarks. Callianassa dijki was described based on isolated major cheliped propodi and a dactylus from the Miocene of Java (Martin, 1883). Later, the species was reported also from the Miocene of Philippines (Martin, 1895; Smith, 1913; Karasawa et al., 2008). Karasawa et al. (2008) reassigned Callianassa dijki to Neocallichirus. Böhm (1922) reported further specimens of C. dijki from the Miocene of Java, and described a new species, Callianassa frangens Böhm, 1922, based on an isolated fragmentary dactylus, coming from the same strata as C. dijki. The morphology of cheliped elements of both species, C. dijki and C. frangens, is very close to K. karumba, the type species of the genus. Callianassa frangens is herein considered a name given to a major cheliped dactylus belonging to C. dijki, and as such it is treated as a junior subjective synonym of the latter species.

Karumballichirus khadroensis ( Hyžný and Charbonnier in Hyžný, Charbonnier, Merle, Lashari, Bartolini and Métais, 2016 )

? “fragments of chelae of an Uca or Cardisoma”.-Stoliczka, 1871: 2, pl. 1 figs 3-10.

Neocallichirus khadroensisHyžný and Charbonnier in Hyžný et al., 2016: 344, figs. 2, 5A3, C2, D3, 6I.

Karumballichirus khadroensis.-Hyžný et al., 2025: 100, fig. 2.

Remarks. Neocallichirus khadroensis has been reported from the Paleocene (Danian) of Pakistan (Hyžný et al., 2016). The species was reassigned to Karumballichirus by Hyžný et al. (2025).

Karumballichirus lakhraensis ( Hyžný and Charbonnier in Hyžný, Charbonnier, Merle, Lashari, Bartolini and Métais, 2016 )

Calliax sp.-Charbonnier et al., 2013: 106, fig. 2A, B.

Neocallichirus lakhraensisHyžný and Charbonnier in Hyžný et al., 2016: 346, figs. 4, 5A2, B2, C3, D2, E2, 6H.

Karumballichirus lakhraensis.-Hyžný et al., 2025: 101, figs. 3, 4.

Remarks. Neocallichirus lakhraensis has been reported from the Paleocene (as Calliax sp.) and Eocene (Ypresian) of Pakistan (Charbonnier et al., 2013; Hyžný et al., 2016; 2025). The species was reassigned to Karumballichirus by Hyžný et al. (2025).

Karumballichirus maximus (A. Milne-Edwards, 1870 )

Callianassa maxima A. Milne-Edwards, 1870: 97, pl. 2 fig. 5.-Dworschak, 2008: 75.-Schweitzer et al., 2010: 35 (list).

Karumballichirus maximus.-Hyžný et al., 2025: 103, fig. 5.

Not Callianassa (Callichirus) maxima.-Kemp, 1915: 252-257, fig. 21, pl. 13 figs. 1-5.- De Man, 1928: 29, 92, 112 (part).- Daniel, 1981: 193-204, pl. 6 figs. 1, 2.

Not Callianassa maxima.- Pillai, 1954: 23-26, figs. 1-5.

Remarks. Callianassa maxima was reported from the Holocene of Thailand, based on several subfossil chelae recovered while digging a canal in Thailand some distance from the sea (A. Milne-Edwards, 1870). The type material of C. maxima, once considered lost (Sakai, 1999: 103), has recently been found, photographed, and figured by Hyžný et al. (2025). Callianassa maxima was described based on more than one specimen, as was clearly mentioned by A. Milne-Edwards (1870), contrary to all subsequent authors who stated that it was described based on a single subfossil chela (Kemp, 1915: 252; Pillai, 1954: 23; Daniel, 1981: 193; Sakai, 1999: 103; Dworschak, 2008: 75).

A. Milne-Edwards’ figures of the chela are virtually identical to those figured from southern India (Daniel, 1981; Kemp, 1915; Pillai, 1954) so it is not surprising that these authors used this species name for a modern species. Only the size (chela twice as long as extant species) and antiquity prevent us from synonymising K. kempi with K. maximus. Callianassa maxima was considered a separate species by Dworschak (2008). The species was reassigned to Karumballichirus by Hyžný et al. (2025).

Karumballichirus pseudoniloticus ( Lőrenthey in Lőrenthey and Beurlen, 1929 ) n. comb.

(Fig. 4)

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Figure 4.
Karumballichirus pseudoniloticus (Lőrenthey in Lőrenthey and Beurlen, 1929) n. comb., Eocene (Lutetian) of Spain. A, left major propodus (MGSB 15995); B, left major merus (MGSB 15991); C, left major claw consisting of articulated dactylus, propodus and carpus (MGSB 15990). All specimens are to the same scale. Scale bar = 10 mm.

Calianassa [sic] pseudoniloticaLőrenthey in Lőrenthey and Beurlen, 1929: 53, pl. 1, figs. 10, 11.

Callianassa pseudonilotica.- Glaessner, 1929: 88.- Vía Boada, 1969: 41, text-fig. 2, pl. 1 figs. 9-15. -Schweitzer et al., 2010: 36.

Remarks. Lőrenthey in Lőrenthey and Beurlen (1929) described the species based on several isolated major cheliped propodi with broken fixed fingers, originating from the middle Eocene (Lutetian) of Hungary. Later, Vía Boada (1969) presented more complete material from the middle Eocene (Lutetian) of Spain, including ischium, merus, carpus, propodus, and dactylus. The species is morphologically very close to modern species of Karumballichirus but lacks pronounced tuberculation on the major propodus. Thus, C. pseudonilotica is reassigned to Karumballichirus.

Karumballichirus pustulatus ( Withers, 1926 ) n. comb.

(Fig. 5)

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Figure 5.
Karumballichirus pustulatus (Withers, 1926) n. comb., ?Eocene-Oligocene of the Scotland Beds, Barbados, isolated major cheliped propodi in lateral (1), mesial (2), and frontal (3) views. A, paratype NHMUK 25778; B, paratype NHMUK 25777; C, paratype NHMUK 25779; D, holotype NHMUK 25774; E, paratype NHMUK 25775; F, paratype NHMUK 25776. All specimens are to the same scale. Scale bar = 10 mm.

Callianassa pustulataWithers, 1926: 106, pl. 9 figs. 5-9.- Glaessner, 1929: 91.- Donovan, 2010: 42.- Schweitzer et al., 2010: 37 (list).

Remarks. From the ?Eocene-Oligocene strata (Scotland Beds) of Barbados, Withers (1926) reported several ghost shrimp species, one of them Callianassa pustulata. All collected specimens represent isolated major propodi. The presence of a quadrant-shaped tooth at the base of the fixed finger suggest attribution to the genus Karumballichirus. The species is reassigned to that genus.

Karumballichirus trechmanni ( Withers, 1924 ) n. comb.

(Fig. 6)

thumbnail of Figure 6.
Figure 6.
Karumballichirus trechmanni (Withers, 1924) n. comb., Eocene of the Yellow Limestone, Jamaica, isolated major cheliped elements. A, propodus (holotype of Callianassa trechmanni NHMUK 23002) in lateral (1) and mesial (2) views; B, propodus (paratype of C. trechmanni NHMUK 23003) in lateral (1), frontal (2), and mesial (3) views; C, propodus (paratype of C. trechmanni NHMUK 23004) in lateral (1), frontal (2), and mesial (3) views; D, propodus (holotype of C. subplana NHMUK 23013) in lateral (1) and mesial (2) views; E, propodus (paratype of C. subplana NHMUK 23014) in lateral (1) and mesial (2) views; F, propodus (NHMUK 60517) in lateral view; G, propodus (NHMUK 60511) in lateral view; H, propodus (holotype of C. gigantea NHMUK 23015) in lateral (1), frontal (2), and mesial (3) views; I, dactylus (NHMUK 23020) in lateral (1) and mesial (2) views; J, dactylus (NHMUK 60528) in lateral view; K, dactylus (NHMUK 23021) in lateral (1) and occlusal (2) views; L, propodus (NHMUK 26008) in lateral (1) and mesial (2) views; M, merus (NHMUK 23023) in lateral (1) and mesial (2) views; N, propodus with articulated dactylus (NHMUK 26009) in lateral view. All specimens are to the same scale. Scale bar = 10 mm.

Callianassa TrechmanniWithers, 1924: 83, pl. 2 figs. 1-8.- Glaessner, 1929: 93.- Vía Boada, 1969: 47, 56.- Morris, 1993: 122, fig. 2.8-9.- Donovan, 2010: 42.-Schweitzer et al., 2010: 37 (list).

Callianassa subplanaWithers, 1924: 85, pl. 2, figs. 9-12.-Glaessner, 1929: 91.- Vía Boada, 1969: 47, 56.- Morris, 1993: 120, figs 2.4-5.- Donovan, 2010: 42.- Schweitzer et al., 2010: 37 (list). (n. syn.)

Callianassa giganteaWithers, 1924: 86, pl. 2, figs. 13, 14.- Glaessner, 1929: 81.- Vía Boada, 1969: 56.- Morris, 1993: 120, figs. 2.6-7.- Donovan, 2010: 43.- Schweitzer et al., 2010: 35 (list). (n. syn.)

Xanthilites rathbunaeWithers, 1924: 89, pl. 4 figs. 5, 6.- Morris, 1993: 122, fig. 1.10.

Not Xanthilites rathbunaeWithers, 1924: 89, pl. 3 figs. 1-6.- Morris, 1993: 122, figs 7-9.

?Varuna sp. Withers, 1924: 90, pl. 4 fig. 7.- Glaessner, 1929: 392.- Morris, 1993: 122, fig. 1.11.- Donovan, 2010: 42.

Remarks. From the Eocene (?Lutetian) strata of Jamaica (the Yellow Limestone), Withers (1924) described three species of Callianassa, based on isolated major cheliped propodi. The re-examination of the type material by one of us (MH) revealed that all three taxa represent a single species, with differences among them being attributable to intraspecific variation. Thus, Callianassa subplana and Callianassa gigantea are considered synonymous with Callianassa trechmanni, as already suggested by Vía Boada (1969: 56). It is of note, that fragmentary fingers identified by Withers (1924) as claw remains of Xanthilites rathbunae, a crab species reported from the same strata, represent dactyli of major chelipeds of C. trechmanni as revised herein (Fig. 5I-K). Moreover, an isolated major cheliped merus (Fig. 5M) was reported by Withers (1924) as questionably belonging to a thoracotrematan crab (as ?Varuna sp.). The correct identification of these elements was already recognized by Vía Boada (1969), but overlooked by Morris (1993). All the material except the chela of C. gigantea, originated from a single locality (Eo1 sensu Donovan, 2010), while a claw of C. gigantea was reported from a different outcrop (locality Eo2 sensu Donovan, 2010); strata at both locations are coeval.

The morphology of propodi, dactyli, and the single merus is very close to extant representatives of Karumballichirus as presented above, including a typical quadrant-shaped tooth at the base of the fixed finger. Thus, C. trechmanni is reassigned to the genus Karumballichirus.

Karumballichirus tuberculatus ( Lőrenthey in Lőrenthey and Beurlen, 1929 )

Calianassa [sic] tuberculataLőrenthey in Lőrenthey and Beurlen, 1929: 51, pl. 1 fig. 9.

Callianassa tuberculata.- Glaessner, 1929: 93.

Neocallichirus borensisBeschin, De Angeli, Checchi and Mietto, 2006: 97, fig. 2, pl. 1 figs. 4-6.- De Angeli and Garassino, 2006: 16.- Schweitzer et al., 2010: 38 (list).- Hyžný et al., 2016: 348, fig. 6A-C.

Neocallichirus tuberculatus.- Hyžný et al., 2016: 350, figs. 6D-F.

Karumballichirus tuberculatus.- Hyžný and Zorn, 2020: 21, pl. 5 figs. 3-5.

Remarks. Callianassa tuberculata was described from the middle Eocene of Hungary based on fragmentary major cheliped elements (propodi and dactyli) (Lőrenthey and Beurlen, 1929). Hyžný and Zorn (2020) considered the species a senior subjective synonym of yet another species, Neocallichirus borensisBeschin, De Angeli, Checchi and Mietto, 2006, originally described from the upper Eocene of Italy (Beschin et al., 2006). Hyžný and Zorn (2020) reported on new occurrences from the Eocene of Austria and Italy, and reassigned the species to the genus Karumballichirus.

Karumballichirus vidali ( Vía Boada, 1959 ) n. comb.

(Fig. 7)

thumbnail of Figure 7.
Figure 7.
Karumballichirus vidali (Vía Boada, 1959) n. comb., Eocene (Ypresian) of Spain; left major cheliped (holotype MGSB 16027). Scale bar = 10 mm.

Callianassa vidaliVía Boada, 1959: 357, text-fig. 1.- Vía Boada, 1969: 48, text-fig. 3, pl. 1 figs. 16-32.- Schweitzer et al., 2010: 37.

Remarks. The species was described from the lower Eocene (Ypresian) strata of Spain based on well-preserved specimens, including articulated major propodi with ischium, merus, carpus, propodus, and dactylus (Vía Boada, 1959; 1969). Based on close morphological affinity with modern representatives of Karumballichirus, the species is reassigned to that genus. Karumballichirus vidali (Vía Boada, 1959) n. comb. is close to its slightly younger congener, K. pseudoniloticus n. comb.

CONCLUSIONS

The probable antiquity of K. maximus and its size argue against applying this species name to a modern species. Sakai (1999) justified the need for a new name for the extant species on size and the “armament of denticles” of his type but the intensity of denticles is highly variable in the material illustrated so far and seen by us. While genetic differences support the recognition of two extant species, their morphological separation is quite subtle. Based on major cheliped material, ten fossil species are recognized.

ACKNOWLEDGEMENTS

We thank the following for access to material: Paul Clark and Claire Mellish, Natural History Museum, London; Tim-Yan Chan, Institute of Marine Biology and Center of Excellence for the Oceans, National Taiwan Ocean University, Taiwan; Andrew Hosie, Western Australian Museum, Perth; Gavin Daley, Northern Territory Museum and Art Gallery; Sebastián Calzada, Museo Geológico del Seminario de Barcelona.

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  • Consent for publication
    All authors declare that they have reviewed the content of the manuscript and gave their consent to submit the document.
  • Funding and grant disclosures
    The research of MH was supported by European Commission´s Research Infrastructure Action via SYNTHESYS Project (GB-TAF 4495).
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  • Data availability
    All study data are included in the article.
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Edited by

  • Editor-in-chief:
    Christopher Tudge
  • Associate Editor:
    Christopher Tudge

Data availability

All study data are included in the article.

Publication Dates

  • Publication in this collection
    17 Apr 2026
  • Date of issue
    2026

History

  • Received
    07 Aug 2025
  • Accepted
    06 Oct 2025
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Sociedade Brasileira de Carcinologia Instituto de Biociências, UNESP, Campus Botucatu, Rua Professor Doutor Antônio Celso Wagner Zanin, 250 , Botucatu, SP, 18618-689 - Botucatu - SP - Brazil
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