Spinicaudata Catalogus Branchiopoda) (Crustacea:

© 2020 Academia Sinica, Taiwan Open Access Special Issue: Fossil and Modern Clam Shrimp (Branchiopoda: Spinicaudata, Laevicaudata) Spinicaudata Catalogus (Crustacea: Branchiopoda) D. Christopher Rogers Kansas Biological Survey, and The Biodiversity Institute, The University of Kansas, Higuchi Hall, 2101 Constant Avenue, Lawrence, KS 66047-3759, USA. E-mail: [email protected] Received 18 October 2019 / Accepted 14 January 2020 / Published 5 August 2020 Special issue (articles 32-46) communicated by Thomas A. Hegna and D. Christopher Rogers The Spinicaudata (spiny clam shrimp) are a large group of freshwater, bivalved branchiopod crustaceans in need of taxonomic revision. Herein, the extant Spinicaudata families and genera are defined and diagnosed according to modern standards. An annotated catalogue of the Spinicaudata taxa is presented with synonyms. More than 747 spinicaudatan taxa are presented, of which 215 are considered valid families, genera and species. Chresonyms are provided for taxa redescribed according to modern standards. It is hoped that this catalogue will provide a basis for further taxonomic revision and phylogenetic work within the Spinicaudata. Key words: Spiny clam shrimp, Diplostraca, Onychocaudata, Checklist, Systematics. Citation: Rogers DC. 2020. Spinicaudata catalogus (Crustacea: Branchiopoda). Zool Stud 59:45. doi:10.6620/ZS.2020.59-45. BACKGROUND The Spinicaudata (spiny clam shrimp) are the second largest group of freshwater dwelling bivalved branchiopod crustaceans after the Cladocera (Brendonck et al. 2008; Rogers 2009) and have the most confused taxonomy of any branchiopod group. This confusion resulted from a combination of: 1) the great plasticity of the few available morphological characters within the group; 2) the tremendous and poorly understood range of intraand interspecific, generic and familial morphological variation; 3) the lack of sexually selected characters and high number of hermaphroditic lineages; 4) poor and inadequate descriptions and type material; 5) a great disparity between the methods and descriptive language used by clam shrimp palaeontologists and biologists, and; 6) the large number of researchers working in isolation and/or focusing only on local taxa when more holistic approaches were needed. This catalogue is the third produced on the Branchiopoda, the first two being on the Anostraca (Rogers 2013) and the Laevicaudata (Rogers and Olesen 2014). This catalogue is patterned in part on the recent catalogs on decapod crustaceans (e.g., Ng et al. 2008, De Grave and Fransen 2011). Spinicaudatan fossils have been reported from as far back as the Devonian (Tasch 1969) and from all continents, with extant forms known to occur on all continents except Antarctica as well as many oceanic islands (Brendonck et al. 2008, Rogers 2009). Modern clam shrimp occur in seasonally astatic aquatic habitats and inland saline pools and lakes (Brendonck et al. 2008; Rogers 2009). The number of spinicaudatan species has historically been in flux, as many species have myriad synonyms, many nomina inquirenda occur, and many new species are described regularly. Some 195 valid species names are recognized here. Endemicity is high, with 41.7% of species known only from the type locality and 27.1% known from ten or fewer localities. This catalog also reflects the many advances in phylogeny made in recent years (e.g., Schwentner et al. 2009 2020a; Weeks et al. 2009), as well as classifications based upon modern genus concepts (e.g., Belk 1989; Rogers et al. 2012). There are 748 taxa presented in this checklist under the suborder Spinicaudata, including four valid families, 16 valid genera, 194 valid species, and 572 synonyms, homonyms, nomina nuda, nomina dubia, species inquirendae, and nomina oblita. Chresonyms Zoological Studies 59:45 (2020) doi:10.6620/ZS.2020.59-45 1 © 2020 Academia Sinica, Taiwan are provided for taxon redescriptions that facilitate identification and evolutionary relationships. Spelling errors (unless widely promulgated) from the literature are not included. Diagnoses of the spinicaudatan higher taxonomic levels are provided. Fossil spinicaudatans are not included and are beyond the scope of this catalogue. It is important to note that clam shrimp bodies and limbs do not preserve well (Tasch 1969), with only the carapaces typically being preserved. Fossil spinicaudatan taxonomy and systematics, especially at the species level is still very much unsettled, and I recommend the excellent work by Astrop and Hegna (2015) as the best possible starting point for understanding the taxonomy and relationships for those organisms, although the morphological phylogeny presented there conflicts with the molecular phylogeny presented in Schwentner et al. (2020a). A Brief History of Spinicaudata Taxonomy Although originally treated with the Laevicaudata and Cyclestherida in the order Conchostraca, morphological and, eventually, molecular studies demonstrated that this concept was invalid (Fryer 1987; Olesen 1998 2000 2007 2009; Negrea et al. 1999; Spears and Abele 2000; Brabrand et al. 2002; deWaard et al. 2006; Stenderup et al. 2006; Richter et al. 2007; Schwentner et al. 2018). The term ‘Conchostraca’ was abandoned as a useful concept in the early 1980s, and the name now conveys no systematic or phylogenetic meaning (Fryer 1987; Olesen 1998 2000; Martin and Davis 2001; Brendonck et al. 2008; Rogers 2009; Ahyong et al. 2011) and should not be used. Following modern methods, all extant clam shrimp groups and the Cladocera are placed within the order Diplostraca, which contains the Laevicaudata (smooth clam shrimp) and the Onchyocaudata (Schwentner et al. 2018). Onchyocaudata comprises the Spinicaudata and the Cladoceromorpha, with Cladoceromorpha comprising Cyclestherida and Cladocera (Olesen 2007 2009; Olesen and Richter 2013). Linnaeus (1761) described the first spinicaudatan clam shrimp: Monoculus lenticularis. Hermann (1804) described a second species, which he called Daphnia gigas. Brongniart (1820) based on material from France, described Limnadia hermanni, giving us the first of the currently recognized genera. All three of these first taxa (and others) were eventually synonymized as Limnadia lenticularis (Linnaeus, 1761). Audouin (1837) erected Cyzicus to contain Limnadia tetracerus Krynicki, 1830 and his Cyzicus bravaisii. Rüppell (in Strauss-Durchheim 1837) erected Estheria for his species E. dahalacensis, with the genus characters given being identical to Audouin’s Cyzicus (Mattox, 1957a). However, Rüppell’s collection was comprised of both Cyzicus and what would eventually be called Leptestheria. Keillhack (1910) recognized that Estheria Rüppell was a homonym of Estheria Robineau-Desvoidy 1830 (Diptera) and thus preoccupied. Joly (1842) in his review of the clam shrimp proposed Isaura to replace Rüppell’s Estheria, rejecting the name Cyzicus. However, Daday (1915) pointed out that Cyzicus had priority and that the name Isaura was debatably preoccupied by Isaures Savingny, 1817 (Cnidaria). Daday (1913a b 1915) moved Estheria dahalacensis Rüppell, 1837 to a new genus: Leptestheria. Bock (1953) argued strongly for maintaining the genus name Isaura, creating a new family for it, ignoring the priority of Cyzicus. Mattox (1957a) officially put the matter to rest by presenting the entire history of the controversy before the IUCN, who put the name Cyzicus on the Official List of Generic Names, and put Estheria Rüppell, 1837 and Isaura Joly, 1842 on the Official List of Rejected and Invalid Names (ICZN 1958). Strangely, Alonso (1996) and Dumont and Negrea (2002) chose to use Isaura over Leptestheria with no explanation. (Some palaeontologists did as well; e.g., Reible 1962). Joly (1842) provided a review of the few European taxa described. The first monographic treatment of Spinicaudata was prepared by Baird (1849), wherein all spinicaudatans were placed in the Limnadiidae. Baird (1849) provides a very interesting history of the discovery of spinicaudatans, describing the great confusion in the taxonomy already apparent in less than 100 years of the group’s taxonomy. Unfortunately, Baird (1849) added to the confusion, by redescribing taxa he had never actually observed and leaving type specimens that were nothing more than dry, empty carapaces. Some of his taxa have subsequently been treated as nomina nuda or inquirenda due to the lack of detail in his descriptions and the condition of some of his type material (Rogers and Padhye 2015). The first real monographs were provided by Daday, wherein he described numerous new and redescribed old species from all over the world (Daday 1913a b 1914 1915 1923 1925 1926). Daday revised the spinicaudatan clam shrimp genera in three papers (Daday 1913a b 1915) creating a certain amount of confusion in the process. The first two were published in separate journals, but both on 12 April. In these two papers he presents new genera, of which Caenestheria and Eocyzicus were nomina nuda, until the descriptions and definitions were published by Daday in 1915. Daday describes Caenestheriella, Eoleptestheria, Leptestheria, and Leptestheriella (in that order) in his 1913a paper, and uses those names in his 1913b paper, but neither paper cites the other. Many of Daday’s taxa page 2 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan and others were defined based on characters of the carapace (number of growth lines, carapace proportions, and ornamentation of the intervals), number of limbs, number of antennomeres, and spine arrangements. Although Daday’s (1913a b 1914 1915 1923 1925 1926) monographs were greatly criticized (e.g., Ueno 1927; Barnard 1929; Brehm 1933; Gauthier 1933; Linder 1945; Botnariuc 1945 1947; Margalef 1953; Straškraba 1965a b 1966) no competing system was developed. Furthermore, although many authors (Vecchi 1922; Gauthier 1933; Linder 1945; Botnariuc 1945 1947; Straškraba 1965a b 1966; Wiltshire 1973; Marinček and Petrov 1985; Petrov and Marinček 1995; Rogers et al. 2012 2017) demonstrated that most traditional characters used to describe spinicaudatans were dependent on the age of the animal or on the nutritive quality of the food received, new taxa were still described using those characters (e.g., Mattox 1953a 1954a b; Nayar and Nair 1968; Hu 1988a), even though many of those authors cited these works and lauded their findings. Brtek (1997 2002) provided the first modern catalogues of all branchiopod taxa, including Spinicaudata. Unfortunately, the text has many problems and has created confusion (criticised in Rogers 2003 2006). The English and editing is poor, and several taxa previously synonymized based on quantified analyses were resurrected without any justification, and little if any mention of the previous analyses. Similarly, two species are shown as valid names simultaneously in two separate genera (Cyzicus crinitus (Thiele, 1900) and C. ellipticus (Sars, 1897) also in Eocyzicus). Naganawa (2001a b) presented a new classification for the Spinicaudata, presenting all large branchiopod crustaceans (Anostraca, Notostraca, and clam shrimp) in a separate subclass from the Cladocera, and furthermore broke up the Spinicaudata into three suborders: Cyclostraca (containing the Cyclestheriidae (of the separate order Cyclestherida)); Spinirostria (containing the Cyzicids and Leptestheriids, divided among five families), and; Procephalida (containing the Limnadiidae, divided among three families). However, none of the previous nor later morphological and molecular work supported Naganawa’s classification. Since 1996, numerous morphological studies (Belk 1996; Martin and Belk 1989; Olesen 1998 2000 2007 2009; Rabet 2010; Orridge 2011; Rabet et al. 2015; Rogers et al. 2017; Schwentner et al. 2012a; Timms 2016a b 2018; Timms and Schwentner 2017; Tippelt and Schwentner 2018), molecular studies using increasingly more powerful analyses (Spears and Abele 2000; Brabrand et al. 2002; de Waard et al. 2006; Hoeh et al. 2006; Stenderup et al. 2006; Richter et al. 2007; Reiger et al. 2010; Sun et al. 2011; Fritsch et al. 2013; Schwentner et al. 2009 2012b 2018 2020a) and combined analyses (Weeks et al. 2009; Schwentner et al. 2011; Rogers et al. 2012; Richter et al. 2007) have resolved the relationships between the Diplostracan suborders, relationships between Spinicaudata families and genera, and informed species definitions. At this time, we have the clearest conceptual understanding of the higher taxonomy and phylogenetic relationships at family level and above that we have ever had. However, there is still much work to be done at genus and species levels. Catalogue structure The catalogue portion follows the format of the recent catalogues on branchiopod (Rogers 2013; Rogers and Olesen 2014) and decapod crustaceans (e.g., Ng et al. 2008; De Grave and Fransen 2011) for taxonomic consistency among these widely used tools. Supraspecific taxa are presented in bold. Genera and species are listed alphabetically. Synonyms are presented following an equal sign (=). Only taxonomically relevant references are included due to space. Where an important analysis is relevant for a synonymy, a reference is provided as “fide” the synonymizer. For example: = Eulimnadia chacoensis Gurney, 1931 fide Martin 1989. Chresonyms are presented parenthetically after the original author and date, and are referred to as “in the sense of” the redescriber. For example: Limnadopsis parvispinus Henry, 1924 sensu Timms, 2009a. Comments are provided as appropriate, including distribution, important type localities, type material locations, and outlier localities. However, many of the determinations here are in need of verification: records are included, but not necessarily verified or substantiated. Historically, most workers only examined taxa from their region, with no comparative analysis against identified material from other areas. To this day, it is a problem despite all the literature that demonstrates that such a myopic view is nearly useless. The Spinicaudata are very plastic, taxonomically confused, and many are poorly described. All information presented here needs to be checked in detail, through additional collections, examination of specimens and if possible molecular studies as well. These errors may be due to my missing a certain piece of literature, or misunderstanding some datum. This catalogue is designed as a starting point for future revisionary work using modern methods and a broad, global perspective of these animals. That being said, I expect that there are errors in this catalogue. page 3 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan RESULTS SYSTEMATICS Spinicaudatan systematics have been problematic from the beginning. However, great strides have been made in the last two decades establishing the families and genera and their evolutionary relationships. One thing is salient: spinicaudatan taxonomic categories must be defined using molecular tools and morphological characters that are informed by good molecular data. At this time, some genera are reasonably well defined, but most need revision. These genera are being used here as categorical groups for species placement, pending proper testing of those genera. I am confident that more genus level clades and species will be revealed. On the other hand, spinicaudatan species are mostly poorly defined, and we are just beginning to understand and appreciate the complexity of the group. The species listed here are nearly all problematic, in need of comparison with related forms, and need detailed molecular study in order to properly revise them. I expect that a great many of the species listed here are invalid. I equally suspect that there are a great many more undescribed species that we have yet to descry even in the material before us. CLASS BRANCHIOPODA Latreille, 1817 SUBCLASS PHYLLOPODA Tasch, 1969 Diplostraca, Gerstaecker, 1866 Spinicaudata Linder, 1945 = Spinirostria Naganawa, 2001b New Combination = Procephalida Naganawa, 2001b New Combination Diagnosis: Branchiopod diplostracan crustaceans with carapace not truly bivlaved, no hinge present. Carapace folded in half longitudinally, generally bearing growth lines. Entire adult animal encompassed within carapace. Antenna I is not subarticulated. Male first two limb pairs modified as claspers to amplex female during mating. Comments: Naganawa (2001b) proposed a major phylogenetic spilt within Spinicaudata, with the Limnadiidae in the “Procephalida” and the remaining spinicaudatan families in the “Spinirostria”. Similarly, Astrop and Hegna (2015) produced a phylogeny hypothesising that the Limnadiidae are the living remnant of the Vertexioidea Kobayashi, 1954, sensu Astrop & Hegna, 2015, and that the remaining extant spinicaudatan families were the living remnant of the Eosestherioidea Zhang & Chen, in Zhang et al., 1976, sensu Chen & Shen, 1985. However, the molecular analyses of Schwentner et al. (2020a) does not support either of these hypotheses. Limnadiidae Burmeister, 1843 = Limniadiidae Burmeister, 1843 nomen nullum fide Tasch 1969 = Limnadiadae Baird, 1849 nomen imperfectum = Limnadidae Girard, 1854 nomen imperfectum = Imnadiidae Botnariuc & Orghidan, 1941 = Estheriinidae (Kobayashi, 1954) = Limnadopseidae Novojilov, 1958 = Limnadopsioidea Novojilov, 1958 = Limnadopsidae Tasch, 1969 = Paraimnadiidae Roessler, 1991a = Metalimnadiidae Roessler, 1995a = Limnadopsinae Dumont & Negrea, 2002 Diagnosis: (modified from Rogers et al. 2012) Cephalic fornicies absent. Rostrum variable, blunt to acute, long or short. Rostrum lacking an apical spine. Compound eyes projecting in ocular tubercle. Frontal organ present, typically pedunculate, sometimes sessile (Metalimnadia and Imnadia). Occipital condyle present or absent. Carapace thin, laterally compressed, umbone present (Limnadopsis), lacking (most genera) or obscure (Metalimnadia). Carapace with or without melanistic pigmentation, growth lines often obscured or absent. Male first two thoracopods with endopod (sensu Olesen 2007) bearing apical suctorial organ or modified tactile setae (absent in Metalimnadia). Eggs 170–250 μm in diameter, varying in shape and ornamentation. Comments: Rogers et al. (2012) conducted a preliminary revision of the limnadiid genera, primarily to create well defined categories for species group revisions. This starting point has allowed for quantitative species revisions, and provided a quantitative basis for describing new taxa (Timms and Schwentner 2012 2017). The eggs are highly ornamented in this family, and the egg morphology is typically species specific. Important works for this family include Belk (1989), Pereira and García (2001), Rogers et al. (2012), and Bellec and Rabet (2016). Bellec and Rabet (2016) report an undescribed genus under the name “Limnadiidae lineage BO sp. 1”, which has been supported in molecular studies (Schwentner et al. 2020a). Australimnadia Timms & Schwentner, 2012 Diagnosis: (modified from Timms and Schwentner 2012 2017). Populations composed of males and hermaphrodites; amplexus has not been observed. Rostrum broadly triangular, with female rostrum more apically rounded. Angle between rostrum and frons from 90° to 100°. Occipital notch, occipital condyle absents. page 4 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Frontal organ pedunculate, length 0.3–2.5x distance of organ from ocular tubercle. Carapace dorsal margin smooth, lacking carinae, hinge line arcuate. Umbone absent. Carapace unpigmented. Muscle scar circular, or angled 35° to 45° from normal, i.e., body horizontal axis. Clasper endopods bearing an apical suctorial organ. Endite IV with apical dense field of long spines. Female IX and X thoracopods with elongated exopod for egg attachment. Thoracic segments with a low medial protrusion or spines. Telsonic ventroposterior angle without spiniform projection. Telson posteriolateral spine rows confluent dorsally, confluence not projecting. Telson spine rows each with 20–25 spines. Telsonic spine rows heteromorphic between portions anterior and posterior of the caudal filaments. Caudal filaments inserted between the eighth and tenth spine pair, on a low or prominent mound. Cercopods sinuate to straight, with a proximal dorsomedial longitudinal row of plumose setae extending 50–80% of the cercopod, and a dorsal cirrus on the apical 10–20% of the cercopod. Eggs double discoid or nearly double discoid with broad facets. Comments: The type species is Australimnadia gigantea Timms & Schwenter, 2012, a junior synonym of Limnadia grobbeni Daday, 1925: 160, by original designation. The eggs of all three species are depicted in Timms and Schwentner (2017). Attributed Species Australimnadia grobbeni (Daday, 1926) = Limnadia grobbeni Daday, 1926 = Austrolimnadia gigantea Timms & Schwentner, 2012 (fide Timms and Schwentner 2017) Comments: Eastern and northern Australia. Australimnadia multifaciata Timms & Schwentner, 2017 Comments: Australia: Western Australia, along the Pilbarra Coast. Australimnadia torqueova Timms & Schwentner, 2017 Comments: Southwestern coast of Western Australia. Calalimnadia Rabet & Rogers, in Rogers et al., 2012 Diagnosis: (modified from Rogers et al. 2012) Hermaphrodites only. Rostrum rounded. Angle between rostrum and frons 100° to 120°. Naupliar eye shape variable from oval to triangular. Occipital notch occipital condyle absents. Pedunculate frontal organ length approximately 1.5x distance of organ from ocular tubercle. Carapace dorsal margin smooth, lacking carinae, hinge line arcuate, rarely sinuate. Carapace surface between growth lines smooth. Umbone absent. Carapace without pigmentation. Muscle scar angle 35° to 40° from normal. Eggs attaching to prolonged exopods of thoracopods IX and X. Thoracic segments smooth or with dorsoposterior ridge margined with spines or setae. Telson with posteriorly directed spiniform projection present on ventroposterior angle, anteriad of cercopod base. Telson posterior margin posteriolateral spine rows dorsally confluent, confluence not projecting. Each row with average of 23 spines. Caudal filament originating between spine rows at third or fifth spines from confluence. Caudal filament never borne on mound. Cercopods straight, elongate, ~3x length of telson ventral margin, each medially with longitudinal row of setae on proximal 80–90%, with apex beyond the cirrus bent dorsally. Setae long and plumose. Setal row terminates with single spine. Cercopod with subapical, dorsal cirrus, extending from 4 to 15% of cercopod length. Egg averages 170 μm in diameter, spherical to subspherical, with broad, round ridges, with narrow slits between ridges. Attributed Species Calalimnadia mahei Rabet & Rogers, in Rogers et al., 2012 = “Undescribed eulimnadoid.” Weeks et al., 2009 = “Undescribed limnadiid.” Hoeh et al., 2006 Comments: Known only from Mauritius Island. Eulimnadia Packard, 1874 = Eulimadia (in error). Sars, 1895, 1896a b = Limnadia Webb & Bell, 1979; Brtek 1997; Naganawa 2001a b = Uenia Naganawa, 2001b Diagnosis: (modified from Rogers et al. 2012) Populations composed of males and hermaphrodites (except E. agassizii which is only composed of hermaphrodites); amplexus is venter to venter. Rostrum variable, blunt to acute, long or short. Angle between rostrum and frons 80° to 100°. Occipital notch occipital condyle absents. Pedunculate frontal organ length approximately 1.55x distance of organ from ocular tubercle. Carapace dorsal margin smooth, lacking carinae, hinge line arcuate, rarely sinuate. Carapace surface between growth lines smooth. Umbone absent. page 5 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Carapace occasionally pigmented. Muscle scar angle from 0° to 90° from normal. Clasper endopods each bearing an apical suctorial organ. Endite IV may be broadly transverse or bear dense apical field of short setae, or a few long setae or spines. Thoracic segments smooth or with dorsoposterior ridge rimmed with spines or setae. Eggs attaching to prolonged exopods of thoracopods VII and VIII or VIII, VIII to IX or XII, IX and X, X and XI, or XI and XII. Telson with a subcercopodal, posteriorly directed spiniform projection on ventroposterior angle, anteriad of cercopod base. Telson posterior margin posteriolateral spine rows confluent dorsally, with confluence not projecting. Each row has from 6 to 22 spines. Caudal filament originating between spine rows at second, third, fourth, fifth, or seventh spines from confluence. Caudal filament borne on projecting mound. Cercopods arcuate, occasionally sinuate. Cercopod with medial longitudinal setal row on proximal 75 to 80%. Setae plumose and long. Setal row terminates with single spine. Cercopod with subapical, dorsal cirrus, extending from 5–30% of cercopod length. Eggs 170–250 μm in diameter. Shape spherical to subspherical or cylindrical to subcylindrical with one end larger than other. Eggs with large rectilinear polygonal depressions separated by ridges, occasionally with lamellar or setaform spines at polygon ridge line confluences (Belk 1989; Martin 1989; Martin and Belk 1989; Rabet 2010). Comments: No type species was designated by Packard (1874). The type for the genus is designated here as Limnadia agassizii. Important works on this genus include Belk (1989), Martin (1989), Martin and Belk (1989), Rabet (2010), Rogers et al. (2012), and Marinone et al. (2016). Species are so far only reliably separated by egg morphology (Belk 1989; Martin and Belk 1989; Rabet 2010; Rogers et al. 2012; Padhye and Kulkarni 2017), including internal characters (Rabet et al. 2012). However, external characters in sediment collected eggs may be affected by the environment (Rabet et al. 2014). Webb and Bell (1979), Brtek (1997) and Naganawa (2001a b) all treated Eulimnadia under Limnadia, however morphological and molecular characters more than justify this genus as distinct (Martin and Belk 1989; Rogers et al. 2012). Reports of undescribed Eulimnadia from the Neotropical region are reviewed in Rogers et al. (2020). Eulimnadia victoriae Brady, 1916 is a Cyclestheria (Cyclestheridia) (fide Brendonck 1999). A single hermaphrodite specimen (lacking eggs) reported from Thailand (Rogers et al. 2012) had a rostral spine. This is the only record of a rostral spine in Limnadiidae, and no other specimens have been found. Attributed Species Eulimnadia acutirostris Daday, 1926 sensu Rabet, 2010; Rabet et al., 2015 = Limnadia acutirostris (Daday, 1926) Comments: Known only from the type locality in either Niger or Mali, in the Niger River Basin. Redescribed by Rabet et al. (2015). Eulimnadia adarensis Rabet & Lluch, in Rabet et al., 2015 Comments: From two pools in the Wagchoodda Region of Mauritania. Eulimnadia aethiopica Daday, 1926 sensu Rabet, 2010; Rabet et al., 2015 = Limnadia aethiopica (Daday, 1926) Comments: The type locality is either in modern day Chad or Cameroon, and is the only known locality for this species. Figured by Monod (1969a) and redescribed according to modern standards by Rabet et al. (2015). Eulimnadia agassizii (Packard, 1874) = Limnadia agassizii Packard, 1874 = Eulimnadia stoningtonensis Berry, 1926, fide Belk, 1989 Comments: USA: New England states. The type locality is Penikese Island, Massachuttesettes. The egg is depicted in Belk (1989). Smith (1992) redescribed the type material and provided SEM images of the egg. Eulimnadia antlei Mackin, 1940 = Limnadia antlei (Mackin, 1940) Comments: USA. The egg is figured in Belk (1989). Eulimnadia astraova Belk, 1989 = Limnadia astraova (Belk, 1989) = Eulimnadia texana in Moore, 1965 (fide Belk 1989) = Eulimnadia inflecta in Moore and Burn, 1969 (fide Belk 1989) Comments: USA. The egg is figured in the original description. Eulimnadia australiensis Timms, 2016a = Eulimnadia australicemsis Timms, 2016a page 6 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan (misspelling) Comments: Australia: northern New South Wales, Queensland. Eulimnadia behningi Smirnov, 1949 = Limnadia behningi (Smirnov, 1949) Comments: Uzbekistan. The egg is unknown. Eulimnadia belki Martin, 1989 = Limnadia belki (Martin, 1989) Comments: ranges from southern México south to northern South America (Rogers and Cruz-Rivera 2020). Brendonck et al. (1990) demonstrates great overlap in the egg morphology among New World taxa with cylindrical eggs. Eulimnadia beverleyae Timms, 2016a Comments: Paroo Desert of New South Wales and Queensland, Australia. Eulimnadia bondi Padhye, Rabet, Kulkarni and Pagni, 2018 Comments: Goa State, India. The eggs are cylindrical. This species should be compared with E. indocylindrova, E. tauluoensis, and E. braueriana. Eulimnadia brasiliensis Sars, 1902 = Limnadia brasiliensis (Sars, 1902) Comments: Brazil, Venezuela (Pereira and García 2001; Rogers et al. 2020). Martin (1989) and Pereira and García (2001) provide images of the egg. Cesar’s (1990) records are actually E. pampa (Marinone et al. 2016). Reible (1962) provides a poor image. Eulimnadia braueriana Ishikawa, 1895 = Limnadia braueriana (Ishikawa, 1895) = Eulimnadia packardiana Ishikawa, 1895 (fide Rabet 2010) = Limnadia packardiana (Ishikawa, 1895) (fide Rabet 2010) = Eulimnadia taoluoensis Hu, 1986a = Limnadia taoluoensis (Hu, 1986a) Comments: Eastern China, Japan, Korea, Taiwan (Hu 1986; Olesen and Grygier 2003; Shen and Huang 2008; Kwon et al. 2010; Rabet 2010; Wang 2014). The type locality for E. braueriana is Kugenuma in modern Kanagawa Province, Japan, and the type locality for E. packardiana is Nikkô in Tochigi Province, Japan. Hu described E. taoluoensis in his 1986a paper, but also presented it as new in his 1986b paper. Naganawa and Orgilijanova (2000) treated E. taoluoensis as a junior synonym of E. braueriana, but provided no explanation. Comparison of the eggs for the two taxa (Shen and Huang 2008: 354, fig. 1D and; Wang et al. 2014: 414, fig. 2B) demonstrate that they are probably synonyms. However, Shen and Huang’s (2008) SEM of the egg from the vicinity (?) of the type locality, depicts eggs that are covered in debris, and were possibly air dried alcoholic specimens (Rogers and Padhye 2015). They appear to be lacking “inflated rims” (Rabet 2010), but this could be an artefact of maturity or preservation and drying. This species needs to be compared with E. indocylindrova and E. bondi. Eulimnadia canalis Timms, 2016a Comments: Australia: northwestern New South Wales and southwestern Queensland. Eulimnadia chacoensis Gurney, 1931 = Limnadia chacoensis (Gurney, 1931) Comments: Paraguay. Brendonck et al. (1990) demonstrates great overlap in the egg morphology among New World taxa with cylindrical eggs. The egg is depicted in Martin and Belk (1989 and Marinone et al. (2016). See discussion in Marinone et al. (2016) regarding E. ovisimilis as a probably synonym of E. chacoensis. Eulimnadia chaperi (Simon, 1886) (fide Padhye & Rabet 2017) = Limnadia chaperi Simon, 1886 = Eulimnadia azisi Subash Babu & Bijoy Nandan, 2010 (fide Padhye and Rabet 2017) Comments: Type locality given is India: Karnataka State: Ballari. Treated as a junior synonym of E. compressa by Daday (1927). Redescribed by Padhye and Rabet (2017). The egg is remarkably similar to E. cryptus, and SEM is required for separation. Eulimnadia colombiensis Sars, 1902 = Limnadia colombiensis Sars, 1902 = Eulimnadia “columbica” Daday unpublished ms name (fide Martin 1989) = Eulimnadia belki Martin, 1989 = Limnadia belki (Martin, 1989) Comments: northern South America (Rogers page 7 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan and Cruz-Rivera 2020). Brendonck et al. (1990) demonstrates great overlap in the egg morphology among New World taxa with cylindrical eggs. The egg is depicted in Roessler (1989 1991b) and in Pereira and García (2001). Eulimnadia contraria Timms, 2016a = Eulimnadia sp. E Schwentner et al., 2015 (fide Timms 2016a) Comments: Central Queensland, Australia. Eulimnadia cryptus Sanoamuang, Padhye, and Rogers, 2020 = Eulimnadia “magdalensis” Rabet, 2010 = Eulimnadia “magdalensis” Padhye & Rabet 2017 Comments: India, Thailand, and Cambodia. The eggs are remarably similar to E. magdalensis of the Americas, and E. chaperi. SEM is required for proper identification. Eulimnadia cylindrova Belk, 1989 = Limnadia cylindrova (Belk, 1989) Comments: From deserts of southern USA and northern México (Rogers and Cruz-Rivera 2020). Brendonck et al. (1990) demonstrates great overlap in the egg morphology among New World taxa with cylindrical eggs. The egg is depicted in the original description and in Pereira and García (2001). Eulimnadia dahli Sars, 1896b (fide Timms 2016a) = Limnadia dahli (Sars, 1896b) Comments: Northern Territory, Queensland, and Western Australia, Australia. The egg is figured by Timms (2016a). Eulimnadia diversa Mattox, 1937 = Limnadia diversa (Mattox, 1937), fide Belk, 1989 = Eulimnadia inflecta Mattox, 1939, fide Belk, 1989 = Eulimnadia thompsoni Mattox, 1939, fide Belk, 1989 = Eulimnadia alineata Mattox, 1953a, fide Belk, 1989 = Eulimnadia ventricosa Mattox, 1953b, fide Belk, 1989 = Eulimnadia oryzae Mattox, 1954a, fide Belk, 1989 = Limnadia oryzae (Mattox, 1954a) = Eulimnadia francesae Mattox, 1953b, fide Belk, 1989 Comments: USA east of the Great Plains, with invasive poulations in California. The egg is figured in Belk (1989). Eulimnadia dubia Daday, 1913a = Limnadia dubia (Daday, 1913a) Comments: New Guinea. The egg is undescribed. Eulimnadia follisimilis (Pereira & García, 2001) Comments: Venezuela. Eulimnadia garretti (Richters, 1882) = Limnadia garretti Richters, 1882 Comments: Tahiti. The egg has not been figured, but this species is being redescribed. Eulimnadia geayi Daday, 1913a = Limnadia geayi (Daday, 1913a) = Eulimnadia “columbica” Daday unpublished ms name (fide Martin 1989) Comments: Mexico to Colombia and Venezuela (Pereira and García 2001; Reed et al. 2015). The egg is depicted by Martin (1989) and Pereira and García (2001). Martin (1989) points out that: “Daday’s (1926) illustration of the egg of E. geayi also shows a spherical egg with somewhat acute surface projections, but this is inconsistent with eggs of E. geayi in the Hungarian Museum. The eggs of E. geayi are short, grooved cylinders with one end of the cylinder slightly wider than the other.” Eulimnadia gibba Sars, 1900 = Limnadia gibba (Sars, 1900) Comments: Tamil Nadu, India. Rogers and Padhye (2015) discuss E. gibba and suggest that it needs closer examination. Eulimnadia gnammophila Timms, 2016a = Eulimnadia dahli in Timms, 2006, Weeks et al., 2006, and Reed et al., 2015 (fide Timms 2016) = Eulimnadia feriensis in Weeks et al., 2006, and in Reed et al., 2015 (fide Timms 2016a) Comments: Occurs across the southern portions of Australia from Western Australia to Victoria. This species is a gnamma (rockpool) specialist. page 8 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Eulimnadia graniticola Rogers, Weeks, & Hoeh, 2010 Comments: Georgia and Florida, USA. Eulimnadia hansoni Timms, 2016a = Eulimnadia sp. G Schwentner et al., 2015 (fide Timms 2016a) = Eulimnadia sp. H Schwentner et al., 2015 (fide Timms 2016a) = Eulimnadia sp. K Schwentner et al., 2015 (fide Timms 2016a) = Eulimnadia sp. O Schwentner et al., 2015 (fide Timms 2016a) Comments: Inland Australia, particularly in the Paroo Desert region. Eulimnadia indocylindrova Durga Prasad & Simhachalam, 2004 (fide Padhye et al. 2015) Comments: India, Thailand (Rogers et al. 2016a). Rogers et al. (2016a) suggest that E. indocylindrova may be a synonym of E. taoluoensis. Images of the egg in Shen and Huang (2008: 354, fig. 1C) are of specimens obscured by debris and were possibly air dried alcoholic specimens before being prepared for SEM study. They appear to be lacking “inflated rims” (Rabet 2010), but this could be an artefact of egg shell maturity or air drying (Rogers et al. 2016a). This species needs to be compared closely with E. braueriana and E. bondi. It is possible that E. braueriana is a senior synonym of E. indocylindrova. Eulimnadia insularis Rogers & Cruz-Rivera, 2020 = Eulimnadia texana (Packard, 1871) in Smith and Wier 1999 Comments: Puerto Rico, Jamaica, Virgin Islands. Brendonck et al. (1990) demonstrates great overlap in the egg morphology among New World taxa with cylindrical eggs. Smith and Wier (1999) present images of the eggs. Eulimnadia kimberleyensis Timms, 2018 Comments: Australia: Western Australia, known only from the Gardner Plateau. This species is a rock pool (gnamma) specialist. Eulimnadia magdalensis Roessler, 1990 sensu Rabet, 2010 Comments: Brazil, Colombia, Venezuela (Roessler 1995a; Pereira and García 2001; Rabet et al. 2012 2014; Godinho et al. 2014; Bellec and Rabet 2016; Marinone et al. 2016; Rogers et al. 2020). Material reported from Cambodia (Rabet 2010; Padhye and Rabet 2017) needs further examination but is probably conspecific with E. cryptus. The egg is pictured in Pereira and García (2001) and Marinone et al. (2016). Eulimnadia margaretae Bond, 1934 sensu Thiéry, 1996, Rabet et al., 2015 = Limnadia margaretae (Bond, 1934) Comments: Oman, United Arab Emirates, Yemmen (Rabet et al. 2015). Redescribed by Rabet et al. (2015). Eulimnadia mauritiana (Guérin, 1837) = Limnadia mauritana (Guérin, 1837) Comments: Mauritius (Simon 1886). Eulimnadia michaeli Nayar & Nair, 1968 sensu Rogers, Dadseepai, & Sanoamuang, 2016a = Limnadia michaeli (Nayar & Nair, 1968) = Eulimnadia khoratensis Rogers, Dadseepai, & Sanoamuang, 2016a Comments: India, Sri Lanka, Thailand (Rogers and Padhye 2015; Rogers et al. 2016a; Padhye and Kulkarni 2017). The types are presumed lost. The eggs are figured in Samyiah et al. (1985), Rogers et al. (2016a), and also in Padhye and Kulkarni (2017), who examine morphological variation in this species. Eulimnadia orinoquiensis (Roessler, 1991b) fide Rogers et al. 2020 = Limnadia orinoquiensis Roessler, 1991b Comments: Colombia. Eulimnadia ovilunata Martin & Belk, 1989 = Eulimnadia sp. A. Martin, 1989 Comments: Argentina, Brazil (Martin and Belk 1989; Marinone et al. 2016). The egg is pictured in Marinone et al. (2016) and in the original description. Eulimandia ovisimilis Martin & Belk, 1989 = Eulimnadia ovismilis (Belk, 1989) in error in Durga Prasad & Simhachalam, 2004 page 9 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Cape York, in northern Queensland, Australia. The type locality is in coastal sand dunes. Paralimnadia centenaria (Timms, 2016a) fide Timms and Rogers, 2020 = Eulimnadia centenaria Timms, 2016a Comments: Australia: Katherine area of Northern Territory. Paralimnadia cygnorum (Dakin, 1914) sensu Timms, 2016b = Limnadia cygnorum Dakin, 1914 Comments: Australia: southern Western Australia. The type locality is given as Cannington on the Swan River. The egg is figured by Timms (2016b). Paralimnadia datsonae (Timms, 2015) fide Timms and Rogers, 2020 = Eulimnadia datsonae Timms, 2015 Comments: Australia: southern Western Australia. Paralimnadia feriensis (Dakin, 1914) sensu Timms, 2015, fide Timms and Rogers, 2020 = Eulimnadia feriensis Dakin, 1914 = Limnadia feriensis (Dakin, 1914) Comments: Western Australia, Australia. Redescribed by Timms (2015). Paralimnadia flavia Timms, 2016b Comments: Extreme northern Western Australia and Northern Territories, Australia. Paralimnadia hyposalina Timms, 2016b Comments: Australia: hyposaline pools in southwestern Western Australia. Paralimnadia laharum Timms, 2018 Comments: Australia: Victoria. Endemic to the Grampian Mountains. This species is a rock pool (gnamma) specialist. Paralimnadia marplesi (Timms & McLay, 2005) fide Timms and Rogers, 2020 = Eulimnadia marplesi Timms & McLay, 2005 Comments: New Zealand. Collected originally in 1962 and not reported since. Paralimnadia minyspinosa Timms & Schwentner, 2020 Comments: Australia: New South Wales. Endemic to Gibralter National Park. This species is a rock pool (gnamma) specialist. Paralimnadia monaro Timms, 2016b Comments: Known only from southern New South Wales, Australia. This species occurs in pools in granitic sands and muddy basalt on the Monaro Plateau. Schwentner et al. (2020b) demonstrate that this species is probably at least two highly endemic species. Paralimnadia montana Timms, 2016b Comments: Australia: northwest New South Wales mountains. Occurs in gnammas and muddy pools on basalt. Paralimnadia multispinosa Timms, 2016b Comments: Known only from the Payne’s Find area in southern Western Australia, Australia. Paralimnadia queenslandicus Timms, 2016b = Paralimnadia sp. A Schwentner et al., 2015 Comments: Queensland and adjacent inland New South Wales, Australia. Paralimnadia rivolensis (Brady, 1886) sensu Timms, 2015 = Eulimnadia rivolensis Brady, 1886 = Limnadia rivolensis (Brady, 1886) = Eulimnadia palustera Timms, 2015 fide Timms and Rogers, 2020 Comments: Australia: South Australia, Tasmania, Victoria, Western Australia (Dakin 1914; Gurney 1927). Records for NSW, QLD, and NT in Spencer and Hall are errors. The type locality is given as Rivoli Bay, South Australia. The egg is figured in Timms (2015 and 2016b). Paralimnadia saxitalis Timms, 2016 Comments: Australia: one location each in southern Northern Territories (Uluru) and northeast New South Wales (Mt Kaputar). This species is a gnamma (rock pool) specialist. page 16 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Paralimnadia sordida (King, 1855) sensu Timms, 2016b = Limnadia sordida King, 1855 = Eulimnadia sordida (King, 1855) = Eulimnadia victoriensis Sayce, 1903 fide Timms, 2016b = Limnadia victoriesnsis (Sayce, 1903) Comments: Coastal portions of New South Wales and Victoria, Australia. The egg is figured by Timms (2016b). Paralimnadia stanleyana (King, 1855) fide Sars, 1896b, sensu Timms, 2016b = Limnadia stanleyana King, 1855 = Eulimnadia stanleyana (King, 1855) Comments: Coastal New South Wales, Australia; other records are likely misidentifications (Timms 2016b). This species is a gnamma (rock pool) specialist on sandstone. The egg is figured and the complex nomenclatural history is discussed by Timms (2016b). Paralimnadia urukhai (Webb & Bell, 1979) sensu Timms & Schwentner, 2020 = Limnadia urukhai Webb & Bell, 1979 = Limnadia upukhai Webb & Bell, 1979 in error in Shen and Huang 2008 Comments: Timms and Schwentner (2020) redescribed this species, pointing out that there are two genetic lineages. Eastern portion of the New South Wales/ Queensland border region, Australia. This species is a rock pool (gnamma) specialist. Schwentner et al. (2020b) demonstrate that this species is probably two or three highly endemic species. Although never specifically mentioned in the original description, it would appear that the describers named this species after the Uruk-hai, a fictional breed of half human, half orc (goblin) from J.R.R. Tolkien’s fantasy books, “The Lord of the Rings”. Paralimnadia vinculuma (Timms, 2015) fide Timms and Rogers, 2020 = Eulimnadia vinculuma Timms, 2015 Comments: Australia: southwestern Western Australia. Paralimnadia westraliensis Timms, 2016b Comments: Australia: Western Australia. Paralimnadia wolterecki (Brehm, 1933) New Combination = Eulimnadia wolterecki (Brehm, 1933) = Limnadia wolterecki Brehm, 1933 Comments: Celebes. Reported only once. Based on the drawings by Brehm (1933), this species appears to have the morphological characteristics of Paralimnadia. Cyzicidae Stebbing, 1910 = Estherianae Packard, 1874 = Estheriidae Sars, 1900 = Caenestheriidae Daday, 1913a: 12 (pro partim) = Isauridae Bock, 1953 = Bairdestheriidae Novojilov, 1954, in part = Straskrabiidae Naganawa, 2001b New Combination Diagnosis: (From Schwentner et al. 2020a). Cephalic fornices extending anteriorly to rostral apex. Rostrum variable, blunt to acute, long or short, generally triangular to subquadrate in lateral view. Rostrum with or without an apical spine. Compound eyes fused medially, sometimes projecting in smoothly arcuate ocular tubercle. Frontal organ sessile. Occipital notch present. Carapace thick, generally rounded. Carapace dorsal margin smooth, lacking carinae, hinge line straight. Carapace with or without pigmentation, growth lines obvious, projecting. Umbone present, projecting well above hinge line. Muscle scar rarely visible. Male first two thoracopods with endopod (sensu Olesen, 2007) lacking an apical suctorial organ or modified tactile setae. If modified setae or spines are present these are never arranged in a transverse apical row of spatulate spines. Telson without a ventroposterior, posteriorly directed spiniform projection. Eggs 110– 170 μm in diameter, spherical and generally lacking ornamentation. Comments: Two genera are recognised here. Daday (1913a: 14) designated Cyzicus as the type genus. Novojilov (1954) created Bairdestheriidae for a large number of fossil genera, among which were Opsipolygrapta and Pseudograpta, and then moved several recent cyzicid species into these genera based on their descriptions. However, the relationships are at best dubious, and no subsequent authors have followed this arrangement. Cyzicus Audouin, 1837 = Estheria Rüppell in Strauss-Durchheim, 1837 (pro partim), nomen praeoccupatum = Isaura Joly, 1842 nomen praeoccupatum = Caenestheriella Daday, 1914: 106, fide Margalef, 1953, fide Straškraba 1965b page 17 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan = Caenestheria Daday, 1914: 53 (pro partim) = Bairdestheria Raymond, 1946 = Opsipolygrapta Novojilov, 1954 (pro partim) Diagnosis: (From Schwentner et al. 2020a). Populations composed of males and females (except C. gynecia which is only composed of hermaphrodites); amplexus is venter to venter. Rostrum subtriangular (usually females) to subquadrate (usually males), depending on age and gender. Angle between rostrum and frons 160° to 180°. Occipital notch either deep and narrow, often closed, very shallow or absent. Occipital condyle conical, subacute, length subequal to basal width. Rostral spine generally absents. Carapace valve length ~1.3x valve breadth (umbone to margin). Carapace growth line intervals smooth or ornamented (scarring from algae often mistaken for ornamentation). Carapace typically dark brown, occasionally black, or with yellow markings, often with setae. Clasper endopod apically unarmed, or with a few setae, apical margin crenulate at most. Endite IV broadly transverse to cylindrical, bearing a dense, apical field of short spiniform setae. Thoracic segments smooth or with a central dorsoposterior projection and/ or set of spines or setae. Eggs attaching to prolonged exopods of thoracopods IX and X. Thoracopod exopods lacking a triangular lamina. Telson posterior margin posteriolateral spine rows confluent dorsally, with confluence not projecting. Each row has from 10 to 30 spines depending on species. Caudal filament originating between spine rows at fifth, sixth, or seventh spines from confluence. Caudal filament borne or not on projecting mound. Cercopods arcuate, occasionally sinuate, or straight with distal fourth to third bent dorsally. Cercopod with medial longitudinal setal row on proximal 40–60%. Setae plumose and either long or short. Setal row terminates with single spine. Cercopod with subapical, dorsal cirrus, extending from 60–40% of cercopod length. Eggs smooth, unornamented. Comments: Limnadia tetracerus Krynicki, 1830 is the type species monotypy (Auduoin 1837). Caenestheriella was treated as a junior synonym based on morphological and developmental grounds by Margelef (1953), Straškraba (1965b), Wiltshire (1973), Forró and Brtek (1984), Sassaman (1995), Smith and Gola (2001) and Orridge (2011). Molecular results support this move (Schwentner et al. 2015 2020a). The character for separating the two genera was the form of the rostrum which was triangular in in Caenestheriella, but quadrate in at least male Cyzicus (Daday, 1913a). However, Wiltshire (1973) demonstrated that this was a matter of development at least in Nearctic species; younger animals have a triangular rostrum and older animals a quadrate rostrum, with both forms sexually reproductive. Tiwari (1966) moved Cyzicus indicus and C. boysii into the fossil genus Baidestheria Raymond, 1946. Baidestheria species are diagnosed as having the carapace intervals bearing radial striae as opposed to punctae. Rogers and Padhye (2015) point out that carapace fine characters may not be diagnostic at genus level, as they are affected by epibiontic algal growth and probably by nutrition. García and Pereira (2003) state that the Cyzicidae has not been reported from South America; Daday (1914) reported a specimen of C. jonesi from southern South America, and two Cyzicus nomina dubia were described from Brazil, both based on empty carapaces, so their actual placement is questionable. Many Cyzicus taxa from Africa and Eurasia may be moved to Ozesthehria upon re-examination. Attributed Species Cyzicus aegyptiacus Daday, 1914: 290 Comments: Described from Cairo, Egypt, and not reported since. This species needs to be compared with C. ehrenbergi, C. crinitus, C. donaciformis, and C. paradoxus. Cyzicus algericus Daday, 1914: 261 Comments: Algeria. Cyzicus belfragei (Packard, 1871) = Estheria belfragei Packard, 1871 = Caenestheriella belfragei (Packard, 1871) Comments: Described from Waco, Texas, USA. Mattox (1957b) reported it from Kansas, Oklahoma and Texas. This species needs to be compared with C. mexicanus, C. gynecia and C. morsei. Donald’s (1989) record from Wood-Buffalo National Park, Alberta, Canada needs to be re-examined. Cyzicus bucheti (Daday, 1913a) = Caenestheriella bucheti Daday, 1913a nomen nudum = Caenestheriella bucheti Daday, 1914: 136 Comments: Morocco (Thiéry 1986, Van den Broeck et al. 2015). Cyzicus californicus (Packard, 1874) = Estheria californicus Packard, 1874 = Cyzicus newcombii (Baird, 1866) = Cyzicus setosus (Pearse, 1912) page 18 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan = Estheria setosa Pearse, 1912 = Caenestheriella setosa (Pearse, 1912) fide Schwentner et al. 2020a Comments: Redescribed by Daday (1914: 249, 324). Central and northern México, western USA north to Oregon and South Dakota (Mattox 1957b; MaedaMartínez et al. 2002). California, USA. Packard’s (1874) description of C. californicus is not useful. The type locality for C. setosa is De Witt, Nebraska, USA. This species needs to be compared with C. elongatus. Cyzicus crinitus (Thiele, 1900) = Estheria crinita Thiele, 1900: 568 = Caenestheriella crinita (Thiele, 1900) = Caenestheria crinita (Thiele, 1900) = Caenestheriella echinata (Thiele, 1900) = Baidestheria crinita (Thiele, 1900) = Baidestheria echinata (Thiele, 1900) = Opsipolygrapta crinita (Thiele, 1900) = Opsipolygrapta echinata (Thiele, 1900) Comments: This species was originally described from a pool in Tanzania, but has not since been reported from that part of Africa. Daday (1915), Gauthier (1939), and Monod (1969b) report this species from Chad (at Koussri, on the Cameroon Border), Niger, and Sudan. However, these additional localities seem oddly disjunct, separated from the Tanzanian locality by the wet tropical zone. This species needs to be compared with C. ehrenbergi, C. donaciformis, C. aegyptiacus, and C. paradoxus. Cyzicus donaciformis (Baird, 1849) = Estheria donaciformis Baird, 1849 = Baidestheria donaciformis (Baird, 1849) = Caenestheriella donaciformis (Daday, 1913b) = Cyzicus echinatus (Daday, 1913b) = Opsipolygrapta echinatus (Daday, 1913b) Comments: Sudan (Simon 1886). Figured by Daday (1914: 180). This species needs to be compared with O. crinitus, C. ehrenbergi, C. aegyptiacus, and C. paradoxus. Cyzicus eductus (Daday, 1913b) = Caenestheriella eductus Daday, 1914: 127 = Caenestheriella deducta Daday, 1914 nomen imperfectum in Vecchi 1922 Comments: Israel, Syria (Thiéry 1996). This species should be compared with O. crinitus, C. tetracerus, C. gihoni, C. hierosolymitana, C. ehrenbergi, C. donaciformis, C. aegyptiacus, and C. paradoxus. Cyzicus ehrenbergi (Daday, 1913b) = Caenestheriella ehrenbergi Daday, 1913b nomen nudum = Caenestheriella ehrenbergi Daday, 1914: 152 = Caenestheriella ehrenbergi var. dimorpha Daday, 1914: 152 = Caenestheriella ehrenbergi var. michaelseni Daday, 1914: 155 = Caenestheriella ehrenbergi var. michaelseni Daday, 1914: 159 = Cyzicus ehrenbergi var. dimorpha Daday, 1914: 155 = Cyzicus dimorphus (Daday, 1913b) = Baidestheria dimorpha (Daday, 1913b) = Baidestheria ehrenbergi (Daday, 1913b) = Caenestheriella ehrenbergi var. fimbriata Brehm, 1935 = Cyzicus ehrenbergi var. fimbriata Brehm, 1935 = Cyzicus fimbriatus (Brehm, 1935) = Cyzicus ehrenbergi var. michaelseni (Daday, 1913b) = Baidestheria michaelseni (Daday, 1913b) = Cyzicus michaelseni (Daday, 1913b) Comments: The type locality is given as Egypt, but Daday (1914) also mentions material from Australia (his form michaelseni), which is certainly an error in labelling or identification. This species needs to be compared with O. crinitus, C. donaciformis, C. aegyptiacus, and C. paradoxus. Cyzicus elongatus Mattox, 1957b Comments: California, USA. This species needs to be compared with C. californicus. Cyzicus gifuensis (Ishikawa, 1895) = Estheria gifuensis Ishikawa, 1895 = Caenestheriella gifuensis (Ishikawa, 1895) Comments: Japan. The type locality is Mino, Gifu Province. Figured by Daday (1914: 125). Cyzicus gihoni (Baird, 1859) = Estheria gihoni Baird, 1859 Comments: Israel, Lebanon (Baird 1859, Simon 1886, Daday 1914: 300). This species should be compared with C. tetracerus, C. grubei, C. hierosolymitana, C. paradoxus, C. ehrenbergi, C. donaciformis, C. aegyptiacus, and O. crinitus. Cyzicus grubei (Simon, 1886) sensu Alonso, 1996 = Estheria grubei Simon, 1886 page 19 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan = Caenestheria syriaca nomen nudum Daday, 1913b = Caenestheria syriaca Daday, 1914: 62 = Caenestheria grubei Daday, 1914: 131 = Eocyzicus syriacus (Daday, 1914) (fide Brtek, 1997) Comments: Mediterranean region (Daday 1913 1914; Alonso 1996; Machado et al. 1999; PerezBote 2004). Alonso (1996) redescribed this species, providing excellent drawings. The type locality for C. grubei is Spain, at Ciudad Real, and Alonso (1996) reports the species as endemic to arid regions of the Iberian Peninsula and the Balearic Islands. Daday (1915) gave records from modern Israel and Syria. This species should be compared with C. gihoni and C. hierosolymitana. Cyzicus gynecius (Mattox, 1950) = Caenestheriella gynecius Mattox, 1950 Comments: Massachusetts, New Jersey, New York, Ohio, and Pennsylvania USA (Schmidt and Kiviat 2007; Smith and Gola 2001; Orridge 2011). Apparently males are absent. This species needs to be compared with C. mexicanus, C. belfragei and C. morsei. The eggs are spiny (Smith and Gola 2001). Cyzicus hierosolymitanus (Fischer, 1860) = Estheria hierosolymitanus Fischer, 1860 = Cyzicus hierosolymitanus var. rollei Daday, 1914: 322 nomen dubium Comments: Israel, Jerusalem (Simon 1886). Should be compared with C. gihoni. Both may be synonyms of C. tetracerus, which Daday (1914) says is very similar. Redescribed by Daday (1914: 312). Daday’s form rollei (1914) was only known from empty carapaces. Cyzicus jonesi (Baird, 1862) = Estheria jonesi Baird, 1862 Comments: Cuba (Baird 1849; Daday 1914: 240), although Daday mentions one collection from “America Meridionalis”, which is basically tropical and southernmost America. Packard (1874) had material given to him without locality data and suggested the specimens came from the southern USA or Central America. The types were deposited in the Berlin Museum. Cyzicus ludhianatus (Battish, 1981) = Caenestheriella ludhianata Battish, 1981 Comments: India: Punjab; reported once. Probably a synonym of C. annandalei (Rogers and Padhye 2015). Cyzicus madagascarica (Daday, 1914) = Caenestheriella madagascarica Daday, 1914 = Pseudograpta madagascarica (Daday, 1914) Comments: Madagascar. See comments under C. ruber. Cyzicus mexicanus (Claus, 1872) = Estheria mexicanus Claus, 1872 = Estheria culdwelli Baird, 1862 (fide Simon, 1886) = Estheria dunkeri Baird, 1862 (fide Packard, 1883) = Estheria clarki Packard, 1874 (fide Simon, 1886) = Cyzicus seurati Daday, 1914: 265 (fide MaedaMartínez et al., 2002) Comments: Central and northern México, central and eastern USA, and Canada in Alberta and Manitoba (Packard 1874; Daday 1914; Mattox 1957b; Wolfe 1982; Maeda-Martínez et al. 2002). Redescribed by Daday (1914: 252). Maeda-Martínez et al. (2002) state that C. seurati is a junior synonym. However, Daday (1914) states that the egg is covered in spines similar to that observed in C. jonsei. The types of C. seurati are in the Paris Museum and the eggs should be compared with those of C. mexicanus and C. californicus. Packard’s E. clarki description is not useful, but material was deposited at the Chicago Museum, and thus is no longer extant. This species needs to be compared with C. belfragei, C. gynecia and C. morsei. Cyzicus morsei (Packard, 1871) = Estheria morsei Packard, 1871 = Caenestheriella morsei (Packard, 1871) Comments: Described originally from Iowa, USA (Packard 1871). Other records come from Oklahoma, Nebraska (Mattox 1957b), and South Dakota (Packard 1874). Daday (1915: 140) provides a figure. This species needs to be compared with C. mexicanus, C. gynecia and C. belfragei. Packard’s description is not useful. Cyzicus nepalensis Uéno, 1967 Comments: Nepal. Uéno (1967) did not designate types, nor state where his material was deposited, but did suggest that his species may be conspecific with C. annandalei. However, the cercopods depicted in the original description appear distinct from those of other Indian species (Rogers and Padhye 2015). page 20 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Cyzicus politus (Baird, 1849) = Estheria polita Baird, 1849 Comments: India. The type locality is given as: “India”. Cyzicus rubra (Daday, 1913b) = Caenestheriella rubra Daday, 1913b nomen nudum = Caenestheriella rubra Daday, 1914: 146 = Caenestheriella rubra var. acanthoporus Brehm, 1958 = Cyzicus ruber var. acanthoporus (Brehm, 1958) Comments: Madagascar. This species needs to be compared closely with C. madagascarica, which Daday separates on differences of the carapace and abdominal dorsal spines. Cyzicus sinensis Hu, 1988b Comments: Described from a pool near Hefei, Anhui Province, China. Possibly a species of Ozestheria. Naganawa and Orgiljanova (2000) treat this species as a synonym of C. gifuensis, without any explanation. Cyzicus tetracerus (Krynicki, 1830) fide Auduoin, 1837 = Limnadia tetracerus Krynicki, 1830 = Estheria tetracera (Krynicki, 1830) = Isaura cycladoides Joly, 1842 = Estheria cycladoides (Joly, 1842) = Isaura tetracera (Krynicki, 1830) = Cyzicus cycladoides (Joly, 1842) = Cyzicus borceai Daday, 1914: 257 = Cyzicus chyzeri Daday, 1913b: 40 = Cyzicus dubiosus Daday, 1913b: 292 = Cyzicus fallax Daday, 1914: 275 = Cyzicus hungaricus Daday, 1913: 25 = Cyzicus intermedius Daday, 1913: 36 = Cyzicus romanus Daday, 1914: 244 = Cyzicus sibericus Daday, 1913b: 296 = Cyzicus simoni Daday, 1914: 305 = Caenestheriella variabilis Daday 1913b: 17, fide Brtek & Thiéry, 1995 = Caenestheriella cyrenaicus Vecchi, 1922 = Cyzicus cyrenaicus (Vecchi, 1922) = Cyzicus ornatus Smirnov, 1932 Comments: The type locality is in the vicinity of Kharkiv (Charkov), Ukraine. Widespread and common: Albania, Algeria, Armenia, Azerbaijan, Czech Republic, Egypt, France, Georgia, Hungary, Italy, Poland, Romania, Russia (east through Siberia and into the arctic circle), Serbia, Spain, Tunisia, Turkey, Ukraine, Uzbekistan; “Central Sahara” (Thiele 1900; Gurney 1909; Daday 1913b 1914; Gauthier 1938; Cottarelli 1971; Šrámek-Hušek et al. 1962; Vekhov 1974; Lebedeva 1982; Stoicescu 2004). Type species of the genus by monotypy (Mattox 1957b). Joly (1842) and Alonso (1996) provide excellent drawings. Daday (1914) states that C. sibericus is intermediate among several other taxa that were subsequently treated as synonyms of C. tetracerus. Daday (1914) reported C. simoni only from one locality in Lebanon (Beirut), but that it was very similar to C. tetracerus. Cyzicus ornatus was reported from Siberia. Cyzicus variabilis was redescribed by Stoicescu (2004) and presented as a valid species. However, it should be re-examined using modern standards. Nomina dubia, nuda, and species inquirendae Cyzicus boysii (Baird, 1849) nomen dubium fide Rogers & Padhye 2015 = Estheria boysii Baird, 1849 = Caenestheriella boysii (Baird, 1849) fide Daday 1914 = Caenestheriella similis (Baird, 1849) fide Daday 1914 = Baidestheria similis (Baird, 1849) = Pseudograpta boysii (Baird, 1849) = Bairdestheria boysii (Baird, 1849) in Tiwari 1996 = Cyzicus similis (Baird, 1849) fide Rogers & Padhye 2015 = Estheria similis Baird, 1849 = Bairdestheria similis (Baird, 1849) = Pseudograpta similis (Baird, 1849) Comments: The type locality is given as “India”. Tiwari (1996) treats C. annandalei and associated synonyms, as well as C. similis all as junior synonyms of C. boysii. However, Rogers and Padhye (2015) reported that the types of C. boysii and C. similis are empty, dried carapaces; useless for determination. Furthermore, Daday (1914) was unable to differentiate between the two forms, and could not separate the types from any other Eurasian form, and treated both taxa as species inquirendae. Baird’s description of these two species is entirely based on carapace characters, giving the type locality for both as “India”. Cyzicus brasiliensis (Baird, 1849) nomen dubium fide Daday 1914: 327 = Estheria brasiliensis Baird, 1849 Comments: “Brazil”. Description based on empty carapace. page 21 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Cyzicus bravaisii Audouin, 1837 nomen nudum fide Todd, 1952 Comments: Audouin mentions this name once, but provides no description or data and does not refer to any specimens. Cyzicus caldwelli (Baird, 1852) nomen dubium fide Daday 1914: 328 = Estheria caldwelli Baird, 1852 Comments: Lake Winnipeg, Canada. Description based on carapace. Cyzicus dallasi (Baird, 1852) nomen dubium fide Daday 1914: 329 = Estheria dallasi Baird, 1852 Comments: “Brazil”. Description based on carapace. Cyzicus gubernator (Klunzinger, 1864) species inquirenda fide Daday 1914 = Limnadia gubernator Klunzinger, 1864 = Caenestheriella gubernator (Klunzinger, 1864) Comments: Described from Egypt (Simon 1886). Daday (1914) states that the description is barely sufficient to place this taxon in Caenestheriella (among the taxa that were eventually moved to Cyzicus). Cyzicus lofti (Baird, 1862) nomen dubium fide Daday 1914: 326 = Estheria lofti Baird, 1862 Comments: Type locality is Bagdad, Iraq. Description based on empty carapaces. Cyzicus melitensis (Baird, 1849) nomen dubium fide Daday 1914: 325 = Estheria melitensis Baird, 1849 Comments: Malta, Sicily (Simon 1886). Description based on empty carapaces. Cyzicus paradoxus (Daday, 1914) nomen dubium = Caenestheriella paradoxa Daday, 1914: 110 = Bairdestheria paradoxa (Daday, 1914) = Baidestheria paradoxa (Daday, 1914) Comments: Description based on juvenile specimens. The type locality is given as the Niger River Valley in western Africa. Monod (1969b) reports one male and three females from Sanga, in southern Mali, but stated the determination was problematic. Barnard (1935) states that this taxon is a juvenile Ozestheria australis. However, O. australis is unknown outside of southern seasonally dry Africa. Cyzicus politus (Baird, 1849) nomen dubium fide Daday 1914: 327 = Estheria polita Baird, 1849 = Eocyzicus politus (Baird, 1849) Comments: Types are empty, dry carapaces and the description is based solely on carapace details. Ozestheria Schwentner & Richter, in Schwentner, Just, & Richter, 2015 = Caenestheria Daday, 1914: 53 (pro partim) = Opsipolygrapta Novojilov 1954 (pro partim) Diagnosis: (modified from Schwentner et al. 2015). Populations composed of males and females; amplexus is venter to venter. Male and female rostrum triangular, rostral spine generally absent (sometimes present in O. australis). Ocular tubercle smoothly arcuate. Angle between rostrum and frons 150° to 170°. Occipital condyle either short and rounded or elongated and subacute. Carapace valve length ~1.5 times valve breadth (hinge to margin). Carapace with or without sculpturing between growth lines (scarring from algae often mistaken for sculpture). Carapace typically dark brown. Male thoracopod I with endopod bearing one or more transverse apical rows of flattened, broadly subtriangular denticles (claw-like scales). Endite IV broadly transverse to cylindrical, bearing a dense, apical field of short spiniform setae. Eggs attaching to prolonged exopods of thoracopods IX and X. Thoracopod exopods lacking a triangular lamina. Posterior trunk segments with several medial dorsoposterior spines per segment. Telson posterior margin posteriolateral spine rows confluent dorsally, with confluence not projecting. Each row with 10 to 30 spines. Caudal filament originating between spine rows at fifth, sixth, or seventh spines from confluence. Caudal filament borne on projecting mound or not. Cercopods sinuate to curved. Cercopod with medial longitudinal setal row on proximal 40–60%. Setae plumose and either long or short. Setal row terminates with single spine. Cercopod with subapical, dorsal cirrus, extending from 40 to 60% of cercopod length. Comments: Ozestheria lutraria (Brady, 1886) is the type species by designation. Until now the genus was thought limited to Australia. Review of material and original descriptions, plus the molecular analyses presented in Schwentner et al. (2020a), reveals that the page 22 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan genus extends into Asia and Africa. Many Cyzicus taxa from Africa and Eurasia may be moved to Ozestheria upon re-examination. Ozestheria packardi appears to be a complex of species (Schwentner et al. 2015). Novojilov (1954) erected Opsipolygrapta designating Caenestheriella packardi as the type. Chen and Shen (1985) list Opsipolygrapta as an invalid name. Attributed Species Ozestheria altus (Shu, Rogers, Chen, & Yang, 2015) New Combination = Cyzicus altus Shu, Rogers, Chen, & Yang, 2015 Comments: Yunnan Province, China. Known only from the type locality. Ozestheria annandalei Daday, 1913b New Combination = Caenestheriella annandalei Daday, 1913b = Cyzicus annandalei (Daday, 1913b) = Baidestheria annandalei (Daday, 1913b) = Caenestheriella roonwalli Tiwari, 1962, fide Tiwari, 1996 = Cyzicus roonwali (Tiwari, 1962), fide Tiwari, 1996. = Caenestheriella misrai Tiwari, 1962, fide Tiwari, 1996 = Cyzicus misrai (Tiwari, 1962) fide Tiwari, 1996 Comments: Temperate regions of northern India (Rogers and Padhye 2015). Figured by Daday (1915: 166) and Tiwari (1962: 184). Ozestheria australis Lovén, 1847 New Combinataion = Caenestheria australis (Lovén, 1847) = Caenestheriella australis (Lovén, 1847) = Baidestheria australis (Lovén, 1847) = Eocyzicus australis (Lovén, 1847) = Estheria elizabethae Sars, 1898a fide Wolf in Daday, 1914 = Baidestheria elizabethae (Sars, 1898a) = Caenestheriella joubini Daday, 1913b nomen nudum, fide Barnard, 1929 = Opsipolygrapta joubini (Daday, 1913b) = Caenestheriella joubini Daday, 1914: 148 = Caenestheriella vidua Daday, 1914: 122, fide Barnard, 1929 Comments: Widespread and very common in Botswana, Lesotho, Namibia, South Africa, Zimbabwe (Sars 1898a b; Gurney 1904; Daday 1914; Barnard 1935; Brehm 1958; Brendonck 1999; Nhiwatiwa et al. 2014; Mabidi et al. 2016; Milne et al. 2020). Figured in Sars (1898a) and Daday (1914: 99, 123, 176). Ozestheria berneyi (Gurney, 1927) = Estheria berneyi Gurney, 1927 = Caenestheria berneyi (Gurney, 1927) = Eocyzicus berneyi (Gurney, 1927) Comments: Australia: Queensland, and adjacent New South Wales and South Australia (Timms and Richter 2002). Ozestheria dictyon (Spencer & Hall, 1896) = Caenestheria dictyon Spencer & Hall, 1896 Comments: Australia: Northern Territory. Known only from the type locality at Palm Creek in the James Range (Timms and Richter 2002). Sayce (1903) suggested that this taxon was a juvenile form of O. lutraria. Ozestheria elliptica (Sars, 1896) = Estheria elliptica Sars, 1896 = Caenestheria elliptica (Sars, 1896) = Cyzicus ellipticus (Sars, 1896) = Eocyzicus ellipticus (Sars, 1896) Comments: Australia: Western Australia. Only known from the type locality, at Roebuck Bay (Timms and Richter 2002). Refigured by Daday (1915: 97). Ozestheria indica (Gurney, 1906) New Combination = Caenestheriella indica Gurney, 1906 = Cyzicus indicus (Gurney, 1906) = Baidestheria indicus (Gurney, 1906) = Opsipolygrapta indica (Gurney, 1906) Comments: Southern India and Sri Lanka. Briefly reviewed in Rogers and Padhye (2015). Figured by Daday (1915: 162). Ozestheria lutraria (Brady, 1886) = Estheria lutraria Brady, 1886 = Caenestheria lutraria (Brady, 1886) = Estheria dictyon Spencer & Hall, 1896 (fide Sayce, 1903) = Cyzicus lutraria (Brady, 1886) = Cyzicus dictyon (Spencer & Hall, 1896) = ?Caenestheria dictyon (Spencer & Hall, 1896) = Eocyzicus lutrarius (Brady, 1886) Comments: Australia: New South Wales, South Australia, Queensland. The type locality for lutraria is at Innaminka, South Australia, near the Queensland page 23 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan border. Originally described from an empty carapace. Figured by Daday (1915: 91). Ozestheria mariae (Olesen & Timms, 2005) = Caenestheriella mariae Olesen & Timms, 2005 Comments: Australia: Western Australia. This is a rock pool (gnamma) specialist. The type locality is Bushfire Rocks near Hyden. Ozestheria packardi (Brady, 1886) = Estheria packardi Brady, 1886 = Cyzicus (Estheria) packardi (Brady, 1886) = Caenestheriella packardi (Brady, 1886) = Cyzicus packardi (Brady, 1886) = Estheria packardi var. typica Spencer & Hall, 1896 = Caenestheriella packardi var. typica (Spencer & Hall, 1896) = Cyzicus packardi var. typica (Spencer & Hall, 1896) = Estheria packardi var. cancellata Spencer & Hall, 1896 = Caenestheriella packardi var. cancellata (Spencer & Hall, 1896) = Cyzicus packardi var. cancellata (Spencer & Hall, 1896) = Estheria packardi var. minor Spencer & Hall, 1896 = Caenestheriella packardi var. minor (Spencer & Hall, 1896) = Cyzicus packardi var. minor (Spencer & Hall, 1896) = Opsipolygrapta packardi (Brady, 1886) = Baidestheria packardi (Brady, 1886) = Baidestheria var. typica (Spencer & Hall, 1896) = Baidestheria var. cancellata (Spencer & Hall, 1896) Comments: Arid and semiarid Australia. The type locality is Lake Bonney, SA, between Adelaide and the New South Wales border. Figured by Daday (1915: 118). Appears to represent a complex of at least 14 species (Schwentner et al. 2015). Ozestheria pellucida Timms, 2018 Comments: Australia: Western Australia. Endemic to the Gardner Plateau. This species is a rock pool (gnamma) specialist. Ozestheria pilosa (Rogers, Thaimuangphol, Saengphan, & Sanoamuang, 2013) = Cyzicus pilosus Rogers, Thaimuangphol, Saengphan, & Sanoamuang, 2013 Comments: Laos, Myanmar, Thailand. Ozestheria rubra (Henry, 1924) = Estheria rubra Henry, 1924 = Caenestheria rubra (Henry, 1924) = Cyzicus rubra (Henry, 1924) Comments: Australia: southern Northern Territory, northern South Australia, and western portions of Queensland and New South Wales (Schwentner et al. 2015). Ozestheria sarsii (Sayce, 1903) = Estheria sarsii Sayce, 1903 = Cyzicus sarsi (Sayce, 1903) = Cyzicus sarsii (Sayce, 1903) = Caenestheria sarsi (Sayce, 1903) = Estheria sarsii (Sayce, 1903) = Eocyzicus sarsii (Sayce, 1903) Comments: Australia: South Australia and Western Australia (Timms and Richter 2002, Schwentner et al. 2015). The type locality is given as Boulder City (near Kalgoorlie). Figured by Daday (1915: 57). No types were designated. Species inquirenda Ozestheria rufa (Dakin, 1914) species inquirenda = Cyzicus (Estheria) rufa Dakin, 1914 = Caenestheria rufa (Dakin, 1914) = Eocyzicus sp. Brtek, 1997 Comments: Australia: Western Australia (Timms and Richter 2002). Based on two females and not collected since. Eocyzicidae Schwentner, Rabet, Richter, Giribet, Padhye, Cart, Bonillo, and Rogers, 2020 = Caenestheriidae Daday, 1913b: 12 (pro partim) = Baikalolkhoniinae Naganawa, 1999 = Baikalolkhoniidae Naganawa, 1999 New Combination Diagnosis: (Modified from Rogers et al. 2017; and Schwentner et al. 2020a). Populations composed of males and females; amplexus is venter to venter. Rostrum typically sexually dimorphic. Rostrum subtriangular (usually females) to subquadrate (usually males) or rounded. Rostrum may or may not be armed with an apical spine (sometimes present in juveniles and rarely adults). Angle between rostrum and frons 170° to page 24 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan 190°. Occipital notch very shallow or absent. Occipital condyle low, rounded or absent, length half or less basal width. Carapace valve length ~1.5 times valve breadth (hinge to margin). Carapace growth line intervals smooth or ornamented (scarring from algae often mistaken for ornamentation). Carapace typically brown, occasionally black, sometimes with marginal setae. Clasper endopod apically with a transverse row of one to a few apical scales bearing a marginal fringe. Endite IV broadly transverse to cylindrical, bearing a dense, apical field of short spiniform setae. Thoracic segments smooth or with a central dorsoposterior projection and/ or set of spines or setae. Eggs attaching to prolonged exopods of thoracopods IX and X. Thoracopod epipods lacking a triangular lamella. Telson posterior margin posteriolateral spine rows confluent dorsally, with confluence not or slightly projecting. Each row has from six to 30 spines depending on species and gender. Females typically have more and smaller spines than males. Caudal filament originating between spine rows at fifth, sixth, or seventh spines from confluence. Caudal filament borne on projecting mound. Cercopods arcuate or straight. Cercopod with a dorsomedial longitudinal row of setae or spines on proximal 40 to 60%. Setae plumose and either long or short. Row terminates with single spine. Cercopod with subapical, dorsal cirrus, extending from 50 to 40% of cercopod length. Eggs smooth or with surface polygons. Comments: Two genera are recognised. Naganawa (2001b) treated Eocyzicus as a junior synonym of Cyzicus, however this is not supported by molecular studies (Schwenter et al. 2009; Schwentner et al. 2020a). Naganawa (1999) created Baikalolkhoniinae to accommodate a new species of cyzicid clam shrimp from Russia. Brtek (2002) elevated that taxon to family status with no explanation or justification. Tiwari (1966) reported a rostral spine in some large adult E. bouvieri. Eocyzicus Daday, 1914: 190 sensu Rogers et al., 2017 = Caenestheria Daday, 1913b nomen nudum pro partim = Eocyzicus Daday, 1913b nomen nudum = Caenestheria Daday, 1914 fide Brtek et al., 1984 Diagnosis: As for the family. Comments: Daday described Eocyzicus in 1913b, but still presented the genus as new, with an updated description in 1914. Daday never designated a type species for the genus. However, the first species he mentions (1913a: 91) is Eocyzicus orientalis Daday, 1914, which was fixed as the type for the genus in Schwentner et al. (2020a). Naganawa (2001b) treats Eocyzicus as a synonym of Cyzicus, but this was generally ignored, and is not supported by molecular data (Schwentner et al. 2015 2020a). Rogers (2017) provided a review of the genus. Attributed Species Eocyzicus argillaquus Timms & Richter, 2009 = Eocyzicus sp. B Timms & Richter, 2002 Comments: Australia: New South Wales, Northern Territory, South Australia, Queensland, and Western Australia. Eocyzicus armatus Tippelt & Schwentner, 2018 = Eocyzicus lineage Z Schwentner et al., 2013 Comments: Australia: New South Wales, Northern Territory, Western Australia. Eocyzicus bouvieri (Daday, 1914: 201) fide Padhye & Rabet, 2017 = Eocyzicus perrieri Daday, 1913b nomen nudum = Eocyzicus perrieri Daday, 1914: 214 = Eocyzicus pellucidus Tiwari, 1962, fide Tiwari, 1996 = Eocyzicus maliricus Qadri & Baqai, 1956, fide Tiwari, 1996 = Eocyzicus acuta Nayar, 1965 nomen dubium fide Tiwari, 1996 = Eocyzicus sp. Karande & Inamdar, 1965 fide Rogers & Padhye, 2015 Comments: Siberian Russia to Pakistan and northern India (Daday 1914; Rogers and Padhye 2015). Redescribed by Padhye and Rabet (2017). Originally reported from Himachal Pradesh, India (Daday 1913b), Daday later (1914: 104) stated in the description that this species is from Russia, specifically Tobolsk (just north of Kazakhstan) and Obdorsk, now called Salekhard, on the Arctic Circle. Padhye and Rabet (2017) re-examined the types. The description of E. acutus based upon juvenile females. It should be pointed out that E. bouvieri is not the same as C. bouvieri. This species should be compared with E. orientalis and E. zugmayeri. Eocyzicus breviantennus Tippelt & Schwentner, 2018 = Eocyzicus lineage S Schwentner et al., 2013 Comments: South Australia, Australia. Known only from the type locality: 26°59'48.9"S, 133°24'55.2"E. page 25 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan figure 2f depicts a small lamina epipoditalis on the female limb I. Tiwari (1966) stated there was no justification to move this taxon to a separate family. Brtek (2002) unaccountably presented this genus in the Cyzicidae, and suggested that this genus may belong in its own family, but provided no rationale or evidence. Naganawa (2001a) presented this species in an Appendix as belonging in an “undescribed independent family”, but provided no explanation, and later (2001b) moved the genus to a new family Sewellestheriidae. Naganawa’s (2001b) diagnosis is in no way exclusive of the Leptestheriidae. The fact that the lamina epipoditalis is present in Tiwari’s own drawing, and that the remaining characters he used to define his genus are not exclusive, Sewellestheria is treated here as a junior synonym of Leptestheria. Attributed Species Leptestheria aethiopica (Daday, 1923) = Leptestheriella aethiopica Daday, 1923: 376 Comments: Eastern Africa from Egypt and Ethiopia, to the Niger River Valley. Should be compared with L. theilei. Leptestheria biswasi Tiwari, 1965 Comments: Rajasthan, India. Needs to be compared with L. jaisalmerensis (Rogers and Padhye 2015). Leptestheria brasiliensis Van Weddingen & Rabet, 2020 Comments: Known only from a few pools in Palmas de Monte Alto municipality, Bahia State, Brazil. Leptestheria brevirostris Barnard, 1924: 227 Comments: Damaraland, east of Otjiwarango, Namibia. Figured by Brendonck (1999). Known only from the type locality Tladi. Leptestheria brevispina García & Pereira, 2003 Comments: Venezuela. Leptestheria caeciliae (Gauthier, 1951) = Leptestheriella caeciliae Gauthier, 1951 Comments: “Station 5 – Poull Bourgou” near Tambacounda, Senegal is the only known locality. Should be closely compared with L. laurentii, which is known from one pool in the same vicinity. The two taxa are separated primarily on carapace ornamentation, and are probably the same species. Leptestheria calcarata (Wolf, in Daday, 1923) = Leptestheriella calcarata Wolf in litteris, in Daday, 1923: 366 Comments: Botswana, Namibia, South Africa, (Barnard 1924 1929; Brendonck 1999). Figured in Barnard (1929) and Brendonck (1999). Leptestheria compleximanus (Packard, 1877) = Eulimnadia compleximanus Packard, 1877 = Estheria compleximanus (Packard, 1877) = Leptestheria pestai Daday, 1923: 296, fide MaedaMartínez et al., 2002 = Leptestheria vanhoeffeni Daday, 1913b, nomen nudum = Leptestheria vanhoffeni Daday, 1923, fide MaedaMartínez et al., 2002 = Leptestheria vanhoeffeni var. variabilis Daday, 1923, fide Maeda-Martínez et al., 2002 Comments: Northern México and the Great Plains and southern deserts of USA (Maeda-Martínez et al. 2002; Martin and Cash-Clark 1994; Rogers and Hann 2016). The type locality is Ellis, Kansas, USA. Gurney’s (1931) record of L. vanhoeffeni from Paraguay is probably an error. Leptestheria cristata García & Pereira, 2003 Comments: Venezuela. Leptestheria dahalacensis (Rüppell, in StrausDrckheim 1837) fide Daday, 1913a = Estheria dahalacensis Rüppell, in StrausDürckheim 1837 = Isaura dahalacensis (Rüppell, in Straus-Dürckheim 1837) = Estheria pesthinensis Brühl, 1860 = Estheria pestensis (in error) = Leptestheria tenuis Sars, 1901 = Leptestheria dives Daday, 1913b: 345 = Leptestheria aegyptiaca Daday, 1923: 333 = Leptestheria dives var. securiformis Botnariuc, 1947 = Leptestheria rotundirostris Daday, 1913: 56 = Leptestheria intermedia Botnariuc, 1947 = Leptestheria xinjiangensis Hu, 1987, fide Naganawa & Orgilijanova, 2000 = Leptestheria saetosa Marinček & Petrov, 1992 page 32 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Comments: Armenia, Austria, Azerbaijan, Belgium, Croatia, Czech Republic, Egypt, Eritrea, Ethiopia, Georgia, Hungary, Iraq, Italy, Macedonia, Moldova, Mongolia, Romania, Russia (southern), Serbia, Sudan, Syria, Turkey, Ukraine (Simon 1886; Thiele 1900; Sars 1901; Daday 1913b 1923; Botnariuc 1947; Šrámek-Hušek et al. 1962; Marinček and Petrov 1985; Brendonck et al. 1989; Scanabissi Sabelli and Tommasini 1990; Miličić and Petrov 2007; Dobrynina 2010). Naganawa and Orgilijanova (2000) lumped L. xinjiangensis here, but without any explanation or evidence of material examined. Straškraba (1966) and Marinček and Petrov (1985 1991a b c) describe some of the variation in this taxon. Leptestheria dumonti Subash Babu & Bijoy Nandan, 2010 Comments: Southern India. Padhye and Ghate (2016) provide differential diagnosis. Leptestheria echinata (Mohammad, 1986) = Leptestheriella echinata Mohammad, 1986 Comments: Iraq, apparently only known from the type locality, east of Baghdad. Should be compared with L. iranica. Separated from that species by spinulae along the carapace and growth line margins. The type is deposited at the British Museum (1984.192). Leptestheria gurneyi Padhye & Ghate, 2016 Comments: Rock pool species from Maharashtra, India, known only from the type locality. Leptestheria heterochaeta Daday, 1923: 293 Comments: Algeria. Should be compared with L. mayeti. Leptestheria inermis (Barnard, 1929: 270) = Leptestheriella inermis Barnard, 1929 Comments: North Cape Province and East Cape Province, South Africa (Mabidi et al. 2016). Should be closely compared to L. rubidgei and L. setosa. Figured by Brendonck (1999). Leptestheria iranica (Uéno, 1967) = Leptestheriella iranica Uéno, 1967 Comments: Iran; known only from the type locality. No types were designated, and the material examined may be lost. Should be compared with L. echinata. Leptestheria jaisalmerensis Tiwari, 1962 (Tiwari, 1996) = Leptestheria longispinosa Nayar, 1965, fide Tiwari, 1969 = Leptestheria biswasi Tiwari, 1965, fide Tiwari, 1969 Comments: Central and northern India. Reviewed by Rogers and Padhye (2015). Should be compared with L. biswasi and L. dumonti. Padhye and Ghate (2016) provide differential diagnosis. Leptestheria kawachiensis Uéno, 1926 = Leptestheria nanjingensis Zhang & Shen, in Zhang et al. 1976, fide Naganawa & Orgilijanova, 2000 Comments: Japan. Originally described from “a shallow rice field at Koya, Tomorogimura, [Kawachi Province], near the south bank of the Yodo River”, in modern day Osaka Province. Figured also in Dong et al. (1982: 12). Naganawa and Orgilijanova (2000) synonymised L. nanjingensis, but without any explanation or evidence of material examined. Leptestheria kunmingensis Shu, Rogers, Chen, & Yang, 2015 Comments: Yunnan, China. Known only from the type locality, which has been destroyed by development. Leptestheria laurentii (Gauthier, 1951) = Leptestheriella laurentii Gauthier, 1951 Comments: Known only from “Station 4 - Poull Koz” near Tambacounda, Senegal. Should be closely compared with L. caeciliae, which is know from only one pool in the same area. The two taxa are separated primarily on carapace ornamentation, and are probably the same species. In the original description, Gauthier (1951) gives a key to the genus for western Africa, but omits this species. Leptestheria mayeti (Simon, 1886) = Estheria mayeti Simon, 1886 = Estheria angulosa Simon, 1886 = Isaura mayeti (Simon, 1886) in Alonso 1996 = Leptestheria lybica Colosi, 1921 = Leptestheria cortieri Daday, 1923: 324, fide Thiéry, 1996 = Leptestheria aff. cortieri in Cottarelli & Mura, 1983 page 33 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Comments: Northern Sahara (Gauthier 1930 1938); Algeria, Balaeric Islands, Egypt, Libya, Mauritania, Morocco, Saudi Arabia, Sudan, Tunisia, Yemen, possibly Sicily (Simon 1886; Gauthier 1929; Alonso 1996; Thiéry 1986 1996; Roux and Thiéry 1988; Samraoui et al. 2006; Rabet et al. 2015; Van den Broeck 2015). Redescribed by Daday (1923: 288) and by Alonso (1996), who provided excellent figures. Gauthier (1938) describes some variation. Leptestheria nobilis Sars, 1900 = Leptestheriella nobilis (Sars, 1900) = Leptestheria hendersoni Sars, 1900, fide Simhachalam & Timms, 2012 = Leptestheriella hendersoni Sars, 1900, fide Simhachalam & Timms, 2012 = Leptestheriella gigas Karande & Inamdar, 1960, fide Simhachalam & Timms, 2012 = Leptestheriella maduraiensis Nayar & Nair, 1968, fide Simhachalam & Timms, 2012 = Leptestheria maduraiensis (Nayar & Nair, 1968), fide Simhachalam & Timms, 2012 Comments: India. Redescribed by Daday (1923: 358). Reviewed in Rogers and Padhye (2015). Should be closely compared with L. simhadrii. Padhye and Ghate (2016) provide differential diagnosis. Leptestheria orientalis Spandl, 1925 Comments: Known from a single collection from Borneo, near Sarawak. Leptestheria rubidgei (Baird, 1862) = Estheria rubidgei Baird, 1862 = Estheria macgillivrai Baird, 1862 = Leptestheria macgillivrayi (Baird, 1862) fide Sars, 1899 = Leptestheria siliqva Sars, 1898b, fide Sars, 1899 = Leptestheria braueri Daday, 1923: 280 fide Barnard, 1929 = Leptestheria gigantea Wolf, in Daday, 1923 fide Barnard, 1929 = Leptestheria siliqva v. gigantea Wolf (in litteris), nomen nudum in Daday, 1923 Comments: Botswana, Lesotho, Namibia, South Africa, Zimbabwe (Barnard 1924 1929; Brendonck 1999; Nhiwatiwa et al. 2014; Mabidi et al. 2016; Milne et al. 2020). Figured in Sars (1898b) and Brendonck (1999). Sars (1899) provides some very good figures. Leptestheria sambharensis (Tiwari, 1966) = Sewellestheria sambharensis Tiwari, 1966 Comments: Known only from the type locality: Sambur Lake, Rajasthan, India. Probably extinct. Brief review in Rogers and Padhye (2015). Leptestheria sarsi (Daday, 1923) fide Padhye & Rabet, 2017 = Leptestheriella sarsi Daday, 1923: 362 Comments: Northern India. Redescribed by Padhye and Rabet (2017). Padhye and Ghate (2016) provide differential diagnosis. Leptestheria serracauda Rogers, Dadseepai, & Sanoamuang, 2016a Comments: Rice paddies in Roi Et Province, Thailand. Known only from the type locality and one other adjacent rice paddy. Leptestheria setosa (Barnard, 1935) = Leptestheriella setosa Barnard, 1935: 489 Comments: Known from four specimens from a single location in the Kalahari Desert of South Africa. It is morphologically intermediate between L. rubidgei (the form calcarata) and L. inermis, and should be compared with those two forms closely. Figured by Brendonck (1999). Leptestheria simhadrii (Simhachalam & Timms, 2012) = Leptestheriella simhadrii Simhachalam & Timms, 2012 Comments: Known only from the vicinity of the type locality: India, Racharla Mandal, Prakasam District, Pool at Racharla (15°28'N, 78°58'E). Despite the great variability of the material examined and overlapping characters with L. nobilis, the authors presented this taxon as new (reviewed in Rogers and Padhye 2015). Leptestheria striatoconcha Barnard, 1924: 227 Comments: Common in Southern Namibia and South Africa, reported also from Zimbabwe (Nhiwatiwa et al. 2014; Mabidi et al. 2016; Milne et al. 2020). Figured by Brendonck (1999). Leptestheria thielei (Daday, 1923) = Leptestheriella thielei Daday, 1923: 370 Comments: Tanzania. Should be compared with L. aethiopica. page 34 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan Leptestheria titicacae Harding, 1940 = Leptestheria tucumanensis Halloy, 1979 fide Rogers et al. (2020) = Straskrabia titicacae (Harding, 1940) fide Rogers et al. (2020) = Brtekia tucumanensis (Halloy, 1979) fide Rogers et al. (2020) Comments: Northern Argentina, Bolivia, and Peru (Rogers et al. 2020). Brtek, in his 1997 catalogue, had the following statement after both L. titicacae and L. tucumanensis: “(the pertinence to this genus is uncertain) – probably gen. nov.” but provided no explanation as to his conclusion. Naganawa (2001b) following Brtek’s (1997) lead, made the statement that he “agrees” with Brtek, after “reconfirming” the original records, and “… the fact that at present I have enough evidence to justify in establishing…” moving these taxa to new two genera Brtekia and Straskrabia, respectively. This move was criticised in Rogers et al. (2020) and the taxonomy revised based on examination of material. Leptestheria venezuelica Daday, 1923: 313 sensu García & Pereira, 2003 Comments: Aruba, Chile, Venezuela (Daday 1923; Belk et al. 2002; García and Pereira 2003; Rogers et al. 2020). Leptestheria villigera Thiele, 1907 = Leptestheriella villigera (Thiele, 1907) Comments: Madagascar. Redescribed by Daday (1923: 381). Nomina dubia and species inquirendae Leptestheria longispinosa Nayar, 1965 Comments: Juveniles, fide Tiwari (1996). Maghrebestheria Thiéry, 1988 Diagnosis: Populations composed of males and females; amplexus is venter to venter. Rostrum may be sexually dimorphic. Rostrum subtriangular (females) to rounded (males). Angle between rostrum and frons 170° to 190°. Occipital notch very shallow, broad. Occipital condyle low, truncated or acute, length half or less basal width. Carapace valve length ~1.5–1.75x valve breadth (umbone to margin). Carapace growth line intervals smooth (scarring from algae often mistaken for ornamentation). Carapace sometimes with marginal setae. Clasper endopod apically with ventral scales and apical transverse row of spatulate spines. Endite IV subcylindrical, bearing a dense, apical field of short spines and scales. Thoracic segments with dorsomedial spines or setae, posterior most segments with a medial projection bearing spines. Eggs attaching to prolonged exopods of thoracopod X through XV. Thoracopod exopods bearing a triangular lamina. Telson posterior margin posteriolateral spine rows confluent dorsally, with confluence not or slightly projecting. Each row has 50+ spines becoming apically setaform in the posterior most pairs. Females have a similar spine arrangement to males. Caudal filament originating between spine rows just posterior to confluence. Caudal filament base borne on low mound. Cercopods straight with apex bent dorsally. Cercopod with a dorsomedial longitudinal row of spines on proximal 95%, becoming longer in distal portion of cercopod. Cercopod without subapical, dorsal cirrus. Eggs smooth, subspherical, ~130 μm. Comments: Naganawa (2001a b) and Brtek (2002) treated Maghrebestheria as a synonym of Leptestheria but provided no further explanation. Attributed Species Maghrebestheria maroccana Thiéry, 1988 sensu Alonso, 1996 = Maghrebestheria maroçana Thiéry, 1985 (in error?) in Thiéry, 1986 nomen nudum = Maghrebestheria maroccana Thiéry, 1986b in Thiéry, 1986 nomen nudum Comments: Morocco, Spain (Thiéry 1986 1988; Alonso 1996; Van den Broeck 2015). Redescribed by Alonso (1996), who contributed excellent figures. Thiéry (1986) provided a distribution map, and listed this species under the names “M. maroçana Thiéry, 1985” and M. maroccana Thiéry, 1986b, some two years before the actual description was published. However, the only citation in that reference for “Thiéry 1985” is the original description of an anostracan, and the citation “Thiéry, 1986b” was the actual description cited as in press, although it was not published until 1988. Acknowledgment: This paper is dedicated to my dear friend and collecting buddy, Brian Victor Timms. Very special thanks to Jennifer Ginsburg and my daughter Hazel L. Rogers for all their help with old Russian locality names and translations from Russian, Latin, Italian, Albanian, and French. Very special thanks to my dear friends Chun-Chieh Wang and Shusen Shu for help translating some of the Chinese texts. Authors’ contributions: The author designed the page 35 of 44Zoological Studies 59:45 (2020) © 2020 Academia Sinica, Taiwan study and wrote the manuscript. Availability of data and materials: Not applicable. Competing interests: The author declares that he has no conflict of interests. Consent for publication: Not applicable. Ethics approval consent to participate: Not applicable. REFERENCES Ahyong ST, Lowry JK, Alonso M, Bamber RN, Boxshall GA, Castro P, Gerken S, Karaman GS, Goy JW, Jones DS, Meland K, Rogers DC, Svavarsson J. 2011. Subphylum Crustacea Brünnich, 1772, pp. 1–237. 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